NOAA Technical Report NMFS 99 February 1991

Marine Flora and Fauna

of the Northeastern United States

Erect Bryozoa

John S. Ryland

Peter J. Hayward

U.S. Department of Commerce
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69. Environmental quality and aquaculture systems: Proceedings of the

thirteenth U.S.-Japan meeting on aquaculture, Mie, Japan, October 24-25,

1984, edited by Carl J. Sindermann. October 1988, 50 p.

70. New and innovative advances in biology/engineering with potential

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85. Marine farming and enhancement: Proceedings of the Fifteenth

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86. Benthic macrofauna and habitat monitoring on the continental shelf

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89. Early-life-history profiles, seasonal abundance, and distribution of

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Samuel H. Gruber, and Toru Taniuchi. July 1990, 518 p.

91. Marine flora and fauna of the northeastern United States, hinodermata: Crinoidea, by Charles G. Messing and John H. Dearborn. August

1990, 30 p.
NOAA Technical Report NMFS 99

Marine Flora and Fauna

of the Northeastern United States

Erect Bryozoa

John S. Ryland

Peter J. Hayward

February 1991

u.s. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

John A. Knauss, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

William W. Fox Jr., Assistant Administrator for Fisheries
Foreword _

This NOAA Technical Report NMFS is part of the subseries "Marine Flora and Fauna

ofthe Eastern United States" (formerly "Marine Flora and Fauna of the Northeastern

United States"), which consists of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants

and animals of the eastern United States. The manuals are published at irregular intervals on as many taxa of the region as there are specialists available to collaborate

in their preparation. These manuals are intended for use by students, biologists,

biological oceanographers, informed laymen, and others wishing to identify coastal

organisms for this region. They can often serve as guides to additional information about

species or groups.

The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates

of the Woods Hole Region," edited by R.I. Smith, and produced in 1964 under the

auspices of the Systematics Ecology Program, Marine Biological Laboratory, Woods

Hole, Massachusetts. Geographic coverage of the "Marine Flora and Fauna of the

Eastern United States" is planned to include organisms from the headwaters of estuaries

seaward to approximately the 200-m depth on the continental shelf from Maine to

Florida, but can vary somewhat with each major taxon and the interests of collaborators.

Whenever possible, representative specimens dealt with in the manuals are deposited

in the reference collections of major museums.

The "Marine Flora and Fauna of the Eastern United States" is being prepared

in collaboration with systematic specialists in the United States and abroad. Each manual

is based primarily on recent and ongoing revisionary systematic research and a fresh

examination of the plants and animals. Each manual, treating a separate major taxon,

includes an introduction, illustrated glossary, uniform originally illustrated keys, annotated checklist (with information, when available, on distribution, habitat, life history,

and related biology), references to the major literature of the group, and a systematic

index.

Manuals are available from the National Technical Information Service, United

States Department ofCommerce, 5285 Port Royal Road, Springfield, VA 22161, either

as a paper copy or microfiche, for a charge. Manuals are not copyrighted, and so can

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available in most major libraries and listed at the end of this manual.

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Text printed on recycled paper

ii
Marine Flora and Fauna

of the Northeastern United States

Erect Bryozoa

J.S. RYLAND and P.J. HAYWARD

School of Biological Sciences

University of Wales

Swansea, SA2 BFF

United Kingdom

ABSTRACT

Forty-nine species of erect Bryozoa from a broad range of Cyclostome, Ctenostome, and

Cheilostome families are described and illustrated, and an artificial dichotomous key is provided

for their identification. In general, the marine bryozoan faunas of the northeastern coasts of the

United States are poorly known; species records are sparse and voucher collections few, and it

is certain that many more species occur in this region than are presently known. The species

described here occur in intertidal, coastal or offshore habitats; some are well known and have

been recorded on numerous previous occasions, others have been only rarely reported, while

a few are known to occur commonly in the north of the region but have yet to be recorded south

of Cape Cod. Some of the species described have not been recorded at all on northeastern coasts

of the United States, but are widely distributed in North Atlantic continental shelf habitats and

perhaps occur in similar parts of the outer shelf of this region. This fauna is thus provisional,

but is intended to stimulate further work on the Bryozoa.

Introduction _

Bryozoans are sessile, colonial invertebrates which form

a familiar and often conspicuous part of the hard substratum epifauna in most marine habitats. They are common from the rocky intertidal to the continental slope and

achieve their greatest abundance and taxonomic diversity

on offshore coarse grounds. Remarkably specialised faunas

are also known from the unconsolidated deposits of the

abyssal deeps. Freshwater and estuarine species are known,

indeed some may be especially abundant in favorable

habitats, but the phylum as a whole is predominantly

marine. Approximately 5000 living species of Bryozoa are

presently recognized, although the current pace ofsystematic and environmental research, particularly in understudied areas of the marine realm, is such that this figure

has probably already been exceeded.

Many bryozoan colonies are rather small and may not

be immediately recognized for what they are, whilst other

species develop substantial multilamellar, nodular, bushlike or even coral-like forms which may constitute the

greater part of the sessile epifauna in certain habitats.

Colony form is thus extremely varied, but in all species

can be related to a common structural plan, being developed through the repeated budding of zooids, the constituent modules of all bryozoan colonies. A practical distinction may be made between encrusting species, which

develop patches, sheets, or uniseriallines of zooids closely

applied to the substratum, and the erect species, in which

the free-growing colony is attached to its substratum by

either a broad encrusting base, a slender column of heavily

calcified zooids, or a narrow stalk of specially modified

zooids. Erect bryozoans adopt a range of colony form, from

solid, coralliform structures, to dense, bushy tufts, or diffuse and delicately branched growths. This distinction cuts

across taxonomic arrangements as each of the three living

orders of marine Bryozoa includes a wide array of encrusting and erect species; yet, it is a useful distinction in

that erect bryozoans are readily recognized and removed

from benthic samples.

The marine bryozoan fauna of the northeastern U nited

States has received surprisingly little attention. The early

faunal records of, for example, Verrill (1879, a-d) have

been supplemented by very few subsequent publications.

The fauna of the Woods Hole region was catalogued by

Osburn (1912), and a corrected checklist was published by
2 NOAA Technical Report NMFS 99 _

funiculus

c

B

calcified

frontal wall

\

operculum

membranous

frontal wa II

"'" parietal

muscles

parietal

muscles

compensation

sac

o

---- calcified body wall

lH--- membranous sac

-+-+-----entosaccal cavity

-+---exosaccal cavity

'1'11~r+-----intestine

retractor

muscle

fun icui us ----fL--'

annular

muscles

dilator

muscles----+V)

pharynx-------~

g izzard ----t---HiI)U

retractor

muscles

longitudinal

parietal

m u sc Ie ----+n

'-----------------_._.._--------------

Figure 1

Autozooid structure in four groups of Bryozoa. (A) Stoloniferous ctenostome (Bowerbankia); (B) Anascan Cheilostome; (C) Ascophoran

cheilostome; (D) Cyclostome.

Rogick (1964). The algal epifauna of Woods Hole was the

subject of a more detailed account by Rogick and Croasdale

(1949), who also included records of bryozoan species from

other parts of the northeast, both to the south and the north
__________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 3

of Cape Cod. More recently Abbott (1973) researched the

ecology of the benthic bryozoans of Long Island Sound.

Type specimens of species described from New England

by Verrill, and others, were reviewed and redescribed by

Maturo and Schopf (1968). However, there is a dearth of

local faunal studies, and the distribution and natural history

of bryozoans throughout this large and potentially interesting region are largely unknown. As a consequence, it

is certain that our dichotomous key underestimates the total

species diversity of the northeastern seaboard. It is hoped,

however, that it will assist biologists in their task of surveying and cataloguing the bryozoan fauna, and that new

research will necessitate its revision in the near future.

General Structure _

Bryozoan colonies consist of replicated series of zooids,

each budded asexually from a predecessor. The founding

zooid, or ancestrula, metamorphoses from the sexually

produced larva; all subsequent zooids are genetically identical, and thus the colony must be regarded as the genetic

individual. The basic unit of the colony is the autozooid,

a functionally independent module equipped with feeding

apparatus, gut, nerve net, and reproductive organs. Modified polymorphic zooids occur commonly in all taxonomic

groups and may play important roles in the formation and

maintenance of the colony. The ground plan of the autozooid approximates to a box (Fig. 1). The end of the box

furthest from the colony origin is morphologically anterior;

in encrusting colonies the surface ofthe autozooid applied

to the substratum is dorsal, and the opposing surface,

through which the feeding tentacles are extended, is ventral. This terminology is confusing when applied to a

colonial animal, and especially so in the case of erect bryozoans. A more practical notation defines the end of the

autozooid furthest from its point of origin as distal, and

the opposing end as proximal. The surface of the autozooid usually applied to the substratum is basal, and the

upper surface, through which the tentacles are extended,

is frontal. In some erect species the frontal and basal surfaces are still distinct; in others the autozooids are arranged

in back-to-back whorls, and the entire periphery of the

branch is by definition frontal, while the axis of the branch

represents the converged basal surfaces of its constituent

autozooids. In the tubular cyclostome bryozoans and the

bottle-shaped ctenostomes the autozooid orifice is terminal

(Fig. 1), and the autozooid can be correctly oriented only

by reference to internal morphology. However, even in

these cases, orientation may be approximated by reference

to colony morphology, and the terms basal, frontal, distal,

and proximal, with practice, can be used to orient all but

the most highly modified species.

Each bryozoan autozooid is provided with an eversible,

bell-shaped lophophore of ciliated tentacles, with the mouth

at its vertex. A U-shaped gut, comprising a short, and often

rather thick, pharynx, and an equally short oesophagus,

followed by a tripartite stomach, occupies the body cavity.

The stomach consists of a rounded cardia (the anterior part

of which may be modified as a muscular crushing gizzard),

followed by a large caecum, which constitutes the loop of

the gut; the pylorus commences the ascending arm of the

gut and is followed by a slender intestine. The anus opens

dorsally at the base of the lophophore, close to the mouth

but outside the tentacle bell. The bryozoan body cavity is

a true coelom, with a peritoneal lining, and contains a

coelomic fluid in which free cells may be observed. A nerve

ganglion is situated between the mouth and the anus, from

which nerves radiate to the lophophore and gut, and to

the muscle system of the autozooid. The lophophore, gut,

and associated musculature are collectively termed the

polypide. This curious convention has practical descriptive value. All of the parts of the polypide have similar

blastogenic origins; moreover, in a majority of bryozoans

these parts of the autozooid are broken down and regenerated at regular intervals. Indeed, "polypide recycling"

is a characteristic feature of the life cycle of many species.

Unresorbed residues of degenerated polypides form dense

"brown bodies" that may be voided by the new polypide,

or that may simply accumulate in the body cavity of the

autozooid. The body wall of the zooid, including all of its

skeletal and cellular components, together with the enclosed

space, is frequently termed the cystid.

Staining of bryozoan zooids for microscopical examination reveals a mesh of mesenchymatous tissue traversing

the cystid, and particularly concentrated around the gut.

This is termed the funiculus; it is thought to be important

in the distribution of metabolites both within and between

zooids. In ctenostome and cheilostome zooids funicular

strands appear to pass through interzooidal communication

pores, and if metabolite transfer is at least part of its function, the funiculus may playa role in colony-wide nutrition.

In the cyclostomes the funiculus is only poorly developed,

does not seem to pass between zooids, and its function

remains a puzzle, except that the testis always seems to

undergo development in association with it.

A flexible portion of the outer body wall is withdrawn

through the orifice of the autozooid on retraction of the

lophophores, forming the sheath which encloses the tentacles. Eversion ofthe lophophore is achieved through means

of a hydrostatic system which varies in structural detail

between the four major morphological types of marine

bryozoans. In the cylindrical autozooids of the stoloniferan

Ctenostomata, such as Bowerbankia (Fig. 1, a), the flexible

body wall is deformed by the contraction of transverse

parietal muscles. As internal hydrostatic pressure thus

rises, the antagonistic longitudinal parietal muscles relax

and the tentacle bell is smoothly everted. In the anascan

Cheilostomata (Fig. 1, b), and also in the encrusting

carnosan Ctenostomata, only the frontal body wall of the
4 NOAA Technical Report NMFS 99 _

autozooid remains flexible; this is depressed by contraction of the parietal muscles, and the process of tentacle

eversion is essentially the same as that seen in Bowerbankia.

In the ascophoran Cheilostomata (Fig. 1, c) the frontal

surface of the autozooid is completely calcified and internal pressure is raised by depressing a membrane or the

floor of a compensation sac, which lies beneath the frontal

wall. Seawater flows into the space, or sac, beneath the

frontal wall, compensating for the decreased volume ofthe

cystid, and the lophophore is everted as usual. The tubular

autozooids of cyclostome bryozoans (Fig. 1, d) are typically

heavily calcified and the frontal surface is reduced to a

proportionately tiny terminal membrane, deformation of

which can have only a minor effect on internal hydrostatic pressure. Instead, in all cyclostomes, tentacle eversion is achieved through the internal redistribution of

pressure. The cyclostome polypide is suspended inside a

thin membranous sac which divides the cavity of the cystid

into two parts, an outer exosaccal cavity and an inner

entosaccal cavity. The membranous sac is attached to

the wall of the cystid at just a few points and distally

fuses with the tentacle sheath which encloses the lophophore; its wall is equipped with numerous annular muscles.

As sphincter muscles closing the orifice and sealing the

tentacle sheath are relaxed, the terminal membrane is

depressed by the contraction of dilator muscles, forcing

coelomic fluid proximally. This is accompanied by contraction, perhaps sequential, of the annular muscles of

the membranous sac, and the lophophore is effectively

squeezed out through the autozooid orifice. In all bryozoans, retraction of the lophophore is extremely rapid,

effected by a powerful pair of retractor muscles arising from

the proximal end of the cystid and inserted around the base

of the tentacle wall.

All bryozoan colonies are hermaphroditic. In many

species the zooids also are hermaphroditic, usually with

a clear tendency to protandry. In others, however, male

and female zooids occur, and there may be a morphological

expression ofsexual dimorphism. Gonads are ductless, differentiate from peritoneal tissue in the proximal part ofthe

cystid, and liberate eggs and sperms into the body cavity.

Eggs are shed through a pore-the coelomopore-situated

dorsally at the base of the lophophore, while sperm release

in those species in which it has been observed, occurs

through pores at the tips of the tentacles. Fertilization has

been observed in very few bryozoan species, but genetic

evidence suggests that, although self-fertilization is known

to be possible, outbreeding is the rule, eggs being fertilized by allosperm, perhaps entrained by colonywide feeding

currents, after their extrusion through the coelomopore.

In a minority of bryozoans the embryo develops into a freeswimming planktotrophic larva which grows and develops

over a period of months before settling and metamorphosing into the ancestrular zooid. In the majority the fertilized egg is brooded, sometimes within the maternal zooid,

or within its tentacle sheath, but most often in a specialised brooding structure-the ooecium, or ovicell-or

in a morphologically distinctive brooding zooid-the

gonozooid, or gonooecium. Generally, brooding structures

are diverse in all bryozoan groups and are often used as

taxonomic characters.

Zooid polymorphism is the most distinctive feature

of the Bryozoa. Zooid polymorphs, or heterozooids, may

be distinguished by modifications of the lophophore: in

sexually dimorphic species, for example, the male lophophore frequently has a reduced number of tentacles, and

in sheet-like encrusting species, groups of autozooids

with asymmetric lophophores mark excurrent chimneys

for colonywide surface currents. However, most frequently polymorphism is expressed in skeletal modifications, correlated with often profound modifications of

soft part morphology, usually with complete loss of the

lophophore. Heterozooids may be specialised to serve

narrowly defined functional requirements, in most cases

still hardly understood, or may be simplified structures

which contribute to the architecture of the colony. The

simplest type of polymorph is the kenozooid, essentially

an empty box, a cystid devoid of polypide. Kenozooids

seem to act as space fillers in some sheet-like encrusting

species, maintaining colony integrity in areas of uneven

substrata; in others, kenozooids are budded in strict, and

invariable, spatial relationship to autozooids. In certain

erect ascophorans, kenozooids form a continuous, compartmentalized sheet over the basal surface of the colony,

presumably supporting and strengthening it. The stolons

of creeping ctenostomes such as Bowerbankia (see Fig. 30),

and the rhizoids of erect species in all groups, are simply

series of cylindrical kenozooids; further, spines, scuta

(specialized spines that overarch the front of the zooid in

some Anasca) and other processes can all be viewed as

modified kenozooids. In every case the zooid morph nature

of the kenozooid can be demonstrated by reference to its

origin, development, and morphology, all are budded from

pre-existing zooids and comprise a body wall, lined with

peritoneum, with a coelomic cavity crossed by funicular

tissue. By the same criteria, most ovicells, in the Cheilostomata, can be shown to be formed partly or wholly from

polymorphic zooids. The most diverse, and often spectacular, types of polymorph are the avicularia and

vibracula found among the cheilostome bryozoans. In

these the cystid may be radically different from that of the

autozooid, or may be similar to it in both size and shape,

but the most profound polymorphic expression is seen in

the form of the operculum (which closes the orifice in

cheilostome zooids) and its musculature. In avicularia the

operculum is modified as a heavily chitinized mandible,

in vibracula it forms a whip-like seta. These heterozooids

occur in a bewildering variety of forms, several types often

represented in a single species, and are consequently of

great taxonomic use.
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 5

Ecology _

The occurrence and distribution of bryozoans are influenced primarily by the availability of suitable substrata.

In favorable habitats the diversity of species, and the

apparent ubiquity of certain species, may suggest that

the distribution of colonies is otherwise random. However,

it has been shown that bryozoan populations and communities are distributed according to their varying tolerances to gross environmental parameters, as well as to

microenvironmental characteristics, and may be further

affected by competitive interactions with other sessile

orgamsms.

Only a minority of bryozoans will tolerate brackish water

conditions. This minority includes a number of Ctenostomata and encrusting anascan Cheilostomata which

are largely restricted to estuaries or to coastal embayments with low, fluctuating salinities. Some of these

species may achieve a considerable biomass on pier

pilings, breakwaters, and outflow pipes. On fully marine

coasts the richest intertidal faunas are found on sheltered

rocky shores. Perhaps as many as 40 species may be

collected on a good shore; some of these will be essentially subtidal species which can be found only on the

lowest spring tides, while others are largely limited to

intertidal habitats. Most notable among the latter are

the algal epiphytes, whose association with algal substrata results from habitat selection by the settling larva,

and whose life cycles are attuned to a degree to those

of the host algae. Erect species of Bryozoa are usually

limited to the lower shore. The cheilostomes Scrupocellaria

and Bugula can be found as pendent tufts in shaded crevices,

beneath overhanging rocks or attached to the stipes of

large, lower shore algae. Together with the cyclostome

Crisia, they may form a dense turf on more exposed shores,

intergrown with hydroids, small sponges, and ascidians.

The circalittoral fringe supports rich assemblages ofsessile

animals, and vertical surfaces may bear diverse communities of both encrusting and erect bryozoans. Offshore

gravels and shell deposits are most usually dominated by

encrusting species, 90% of which will be cheilostomes, but

stable bottoms of bedrock or boulders, particularly in

current-swept areas, will support communities of erect

species.

Bryozoan ecology is still an underdeveloped discipline

and much careful work needs to be done on the distribution of species, and species communities, in relation to

environmental factors before the habitat requirements

of individual species begin to be understood. In this respect, the work of Harmelin (1976) is a model of its

kind. The most comprehensive review of bryozoan ecology

is that of Ryland (1976). McKinney and Jackson (1989),

in their significant volume Bryozoan Evolution, review the

evolutionary ecology of bryozoans and discuss the adaptive significance of different colony growth forms.

Geographical Distribution _

Patterns of geographical distribution are known with accuracy for very few marine bryozoan species. In a few particularly well studied areas of the northeast Atlantic region,

a few species may be mapped with confidence; elsewhere

persistent misidentification, taxonomic confusion, and a

dearth of reliable records preclude any statements regarding the geographical distribution of bryozoans. In the past

many species have been regarded as practically cosmopolitan. The names of essentially northeast Atlantic species

have been applied to different species along the whole of

the eastern and western seaboards of North America, and

to Australasian, South American, and even Antarctic

species. A great deal oftaxonomic research is still required

to elucidate the resulting tangle, and an important priority is the compilation of accurate local faunas.

Along the eastern coasts of the United States significant

marine faunal boundaries are known to exist in the region

of Cape Hatteras and at Cape Cod. Maturo (1968) conducted a zoogeographical study of bryozoan faunas of the

southeastern seaboard and showed that their geographical

distributions were in accordance with those of other marine

invertebrates. Only 12 % of a total of 246 bryozoan species

were distributed both north and south of the Cape Hatteras boundary, and a further 8% appeared to be restricted

to the Virginian province, between Cape Hatteras and

Cape Cod. There has been no such survey of northeastern

faunas, and the scale of difference between the bryozoan

faunas ofthe Virginian province and those ofsubarctic and

Arctic Canada remains to be established. Many Arctic and

subarctic species have circumpolar, or at least amphiatlantic distributions. For many of the latter, the Gulf

of the St. Lawrence seems to be a significant southern

boundary, although some possibly extend further south

along the outer continental shelf.

Techniques _

All calcified bryozoans should be fixed in 5% buffered

seawater formalin, and stored in 70 % alcohol. The noncalcified Ctenostomata are best narcotized by sprinkling

menthol crystals onto the surface of bowls offresh seawater,

at ambient temperature; they should be fixed after about

12 hours and stored in 5% seawater formalin. It is often

best to identify ctenostomes alive, when such features as

the extended lophophore and embryo coloration are most

easily seen. Identification of cheilostomes and cyclostomes

requires attention to the finer details of zooid morphology;

a good dissecting microscope, capable of x 50- x 100

magnification, is essential, because very few species may

be accurately identified by characters such as colony form.

It is often best to bleach a small part of the specimen, using

a diluted Chlorox or domestic household bleach; rinse it
6 NOAA Technical Report NMFS 99 _

well In fresh water and dry it, before examining it on

a black background. Cardboard cavity slides of the type

used by paleontologists are ideal for this purpose, and

may be used for the storage of voucher specimens. The

cleaned skeletal structures are readily visible, particularly

if reflection of incident light is reduced by staining the

specimen lightly with washable blue ink or food coloring.

A fine camel hair brush is indispensable for removing

detritus which may obscure important morphological

details.

Glossary _

Adventitious avicularium. One occupying some position on the external wall of an autozooid (if. vicarious).

Ancestrula. First-formed zooid of a colony, derived by

metamorphosis of a free-swimming larva.

Ascus. Sac-like hydrostatic organ in ascophoran Cheilostomata (syn. compensation sac, compensatrix).

Autozooid. The feeding zooid of a bryozoan colony.

Avicularium. Specialized cheilostome zooid with reduced polypide but strong muscles operating a mandiblelike operculum (see adventitious, interzooidal, vicarious).

Basal. Under or reverse side of an encrusting or erect

zooid or colony. Morphologically dorsal (if. proximal).

Brown body. Colored spheroid seen in zooids, resulting

from aggregation of non-histolyzable residue of a degenerated polypide.

Cauda. Thread-like proximal portion of a club-shaped

autozooid.

Coelomopore. Pore in the body wall connecting the

coelom with the exterior; specifically, the pore at the base

of the tentacles through which eggs are shed (syn.

supraneural pore).

Condyles. A pair of oppositely placed processes on which

the operculum pivots in the orifice of ascophoran cheilostomes.

Cystid. Cellular plus skeletal layers of the zooid wall (if.

polypide, zooecium).

Distal. Pertaining to the side away from the ancestrula

or origin of growth.

Frontal. Pertaining to the exposed or orifice-bearing side

of a zooid or colony. Morphologically ventral.

Frontal wall. A calcareous frontal body wall (covering

the ascus in some ascophoran cheilostomes).

Funiculus. Mesenchymatous strands connecting the

polypide with the zooidal wall (actually with the communication pores in the wall, in Cheilostomata and Ctenostomata).

Gonozooid. Zooid modified as a brood chamber.

Heterozooid. A specialised zooid, not an autozooid.

Interzooidal avicularium. One which extends to the

basal surface of the colony, but is wedged in between

autozooids rather than replacing one of them in a normal series.

Kenozooid. Heterozooid without a polypide, and usually

without either orifice or muscles.

Lophophore. The ring of hollow, ciliated tentacles surrounding the mouth of the zooid.

Lyrula. Median tooth, often anvil-shaped, on the proximal side of the orifice in some Cheilostomata.

Mandible. Articulated part of an avicularium, moved

by muscles, and homologous with the operculum of an

autozooid.

Ooecium (pI. ooecia). Ovicell or brood chamber in

Cheilostomata.

Ooeciostome. In Cyclostomata, the opening of the

gonozooid, through which larvae are released. Typically

different in shape from the autozooid aperture, and

usually characteristic for each species.

Operculum. A generally uncalcified lamina, hinged or

pivoting on condyles, which closes the zooidal orifice in

Cheilostomata.

Opesia. In anascan Cheilostomata, the opening below

the frontal membrane which remains after formation of

the cryptocyst.

Orifice. Not the mouth, but the opening in the zooid

body wall through which the lophophore is extended.

In cheilostomes this may be surrounded by a peristome,

the opening at the upper end of which is then called the

secondary orifice.

Ovicell. A globular brood chamber in Cheilostomata

(syn. ooecium).

Peristome. In Cheilostomata, a rim, often elevated or

tubular, surrounding the primary orifice. In Ctenostomata used loosely to describe the cylindrical protuberance of the body wall on which the primary orifice

is situated. In Cyclostomata, the distal tubular portion

of the autozooid.

Polypide. The organs and tissues of a zooid which

undergo periodic replacement; viz., tentacles, tentacle

sheath, gut, musculature and nerve ganglion.

Proximal. Pertaining to the side toward the ancestrula

or origin of growth.

Pseudopore. Tissue filled lacuna in the calcification of

the outer zooid wall in many Bryozoa.

Quincuncial. Arrangement of five objects such that four

are placed at the corners of a rectangle with the other

in the center.

Rostrum. (1) spike-like prolongation of an avicularium;

(2) distal part of avicularium occupied by mandible.

Sinus. Slit at proximal edge of orifice in some ascophoran

Cheilostomata.

Transverse parietal muscles. Short muscles inserted on

the body wall of ctenostomes, the frontal membrane of

anascans or the ascus of ascophorans, which contract to

raise internal hydrostatic pressure.

Vibraculum. In Cheilostomata, a heterozooid with an

operculum modified as a long seta slung between

condyles.

Vicarious avicularium. One that replaces an autozooid . .

In a senes.

Zooid. Single unit of a bryozoan colony. Various types

are distinguished by prefixes.
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 7

Key to Genera and Species _

1 Colony uncalcified, forming fleshy, fingerlike growths, cylindrical, branched or lobate, smooth, or with

a coarse, earthy surface; or creeping stolons, bearing upright clumps of bottle shaped zooids; or stiff, branching tufts with the zooids grouped in spiraled, panpipe series. Zooids simple, never bearing spines or

ovicells (Ctenostomata) 2

Colony calcified, forming thick, branching coralliform structures; or slender, branching series of cylindrical

zooids; or stiff, fan-like growths of lightly calcified zooids. Rigid, or jointed, or frondose, and often quite

flexible, but always calcified to a greater or lesser extent. Zooids often with spines, inflated brooding

structures or various other types of polymorph ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6

2 Zooids arranged singly or in clumps along a slender, stiff or flaccid stolon, or packed together in loose clumps

joined basally by thin stoloniform processes. Colony stiffly erect, or loosely creeping 3

Zooids packed closely together, without stolons, the colony developing a solid, cylindrical, branching or

lobate form 5

3 Zooids elongate, slender, up to 3 mm long; packed together in loose, erect clumps; tapered proximally,

and linked by thin, stoloniform, proximal processes (see Fig. 27) Nolella stipata

Zooids squat, bottle shaped, up to 1 mm long; arranged singly, in groups, or in regular, panpipe-like series

along a slender stolon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4 Zooids in paired, closely packed rows (each resembling a set of panpipes), partly spiralled around a stiff,

erect, branching stolon (see Fig. 29 for Amathia vidovici) Amathia

Zooids single or in loosely ordered clumps along a creeping or dependent stolon. Sometimes developing

dense, tufted growths Bowerbankia (Fig. 2)

Figure 2

Bowerbankia. Scale: 10 mm.

5 Colony smooth surfaced, rubbery to touch, light brown to deep mahogany in color; cylindrical or vermiform,

irregularly branched, or lobate. Attached by a slender stem. Subtidal, but frequently cast up along

strandlines (see Fig. 26 for Alcyonidium verrillI) Alcyonidium

Colony developing dense grey-brown tufts up to 200 mm high, with a coarse, muddy surface; branching

regularly, resembling a small specimen of the green alga Codium. Zooids elongate, the orifices opening at

the tip of each branch (see Fig. 28) Anguinella palmata
8 NOAA Technical Report NMFS 99 _

6 Zooids cylindrical, entirely calcified, with a simple, round terminal orifice; no operculum. Calcification

dense, speckled with opaque white spots, but without regular arrangement of pores. Embryos brooded

in conspicuous, inflated, densely stippled brood chambers. No other polymorphs present. Colony thick

and coralliform, or slender and branching, or distinctly jointed (Cyclostomata) 7

Zooids not as described. Orifice subterminal, with an operculum indicated by a thickened, semicircular

marginal sclerite, or a more distinct chitinous flap. Embryos brooded in ovicells, each borne by a single

autozooid. Colony form various (Cheilostomata) 12

7 Colony slender, jointed. Internodes biserial, with zooid orifices all opening on a defined frontal surface.

Gonozooid inflated, replacing a single zooid in a normal linear series (Crisia)* 8

Colony unjointed and rigid 9

8* Internodes very long, wide and sinuous, commonly consisting of about 18-20 zooids (extremes 10-26).

Gonozooid generally occupying position 5 or E(extremes 3-9) in the internode, strongly projecting above,

its orifice tube short, flared, with transversely elongate, upward facing opening (see Fig. 20) .. Crisia cribraria

Internodes short, commonly consisting of 5-7 zooids (extremes 4-10). Gonozooid generally occupying

position 2 or 3 in the internode, pear-shaped; orifice tube curved forwards, narrowing towards its opening

which is transversely oval and faces obliquely upwards and forwards (see Fig. 21) Crisia sp.

9 Colony consisting of an encrusting portion, ·:omprising linear biserial to multiserial groups of zooids,

giving rise to stout clavate or branching growths, often detached, with zooids arranged in whorls opening

on all faces of the branch (see Fig. 23 for Entalophoroecia deflexa) Entalophoroecia

Colony slender or stout, attached by a featureless encrusting base. Zooids opening on a defined frontal

surface 10

10 Colony bifurcating regularly, each bifurcation assuming an elegant tuning-fork shape. Zooid orifices

arranged in graduated horizontal rows directed alternately to left and right of branch axis, and projecting conspicuously from surface (see Fig. 22) Idmidronea atlantica

Colony stout, with a thickened, rugose base, the branches dividing at irregular intervals, tapering distally,

developing a three-dimensional coral-like structure up to 60 mm high. Zooid orifices projecting slightly

from colony surface at branch tips, but becoming immersed and opening flush with surface 11

11 Basal and frontal surfaces of colony coarsely granular; with large pores and conspicuous longitudinal

ridges and furrows between orifices, and particularly well developed on basal surface (see Fig. 24) ..

· Hornera lichenoides

Basal and frontal surfaces of colony smooth, finely punctate, with a few large pores (see Fig. 25) ...

· . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegohornera violacea

12 Colony flexible; frondose and lightly calcified; or in the form ofshort narrow cylinders linked by chitinous

joints; or jointed or unjointed, freely branching tufts, the zooids in uniserial, biserial or multiserial

arrangements, but always lightly calcified, and with part of their frontal surface membranous .

· (Anasca + Haplota) 13

Colony rigid, non-flexible, without joints. Slender, robust, or even nodular, but always heavily calcified;

no membranous area on the frontal surfaces of zooids, although the calcified wall may be overlain by

a thick, translucent cuticle (Ascophora) 41

13 Colony frondose; fronds broad or narrow but multiserial, (i.e., composed of many parallel series of zooids),

bilaminar (i.e., of two layers of zooids placed back to back) or unilaminar 14

• The available literature suggests that other imperfectly characterized species of Crisia occur in the area covered by this manual. Descriptions of North

Atlantic Crisiidae are given by Harmer 1891; Kluge 1962, 1975; Ryland 1967; and Hayward and Ryland 1978, 1985.
__________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 9

Colony form variable, but fronds or branches always narrow, and never both multiserial and bilaminar 16

14 Colony unilaminar, translucent. Zooids without spines. No avicularia or ovicells .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carbasea carbasea (Fig. 3)

Colony bilaminar. Zooids with or without spines. Both avicularia and ovicells occur 15

Figure 3

Carbasea carbasea. Scale: 20 mm.

15 Fronds broad or palmate, the lobes terminally rounded; zooids with two to six spines distally; avicularia

placed at the bifurcation of zooid rows Flustrafoliacea (Fig. 4)

Fronds slender, the lobes wedge shaped with squared ends; zooids without distal spines; avicularia placed

in the zooid rows, not at bifurcations Securiflustra securifrons (Fig. 5)

Figure 4

Flustra foliacea. Scale: 20 mm.

Figure 5

Securiflustra securifrons. Scale: 20 mm.
10 NOAA Technical Report NMFS 99 _

16 Tall (to 10 cm) whitish colonies; fronds slender, composed of paired zooids placed back to back in single,

branching series Eucratea loricata (Fig. 6)

Colony form variable; fronds uni- or multi-serial, but never composed of zooid pairs placed back to

back 17

Figure 6

Eucratea loricata. Scale: 20 mm.

Figure 7

Celiariafistulosa. Scale: 2.5 mm.

17 Colony jointed; branches cylindrical, dividing dichotomously at the joints; zooids in longitudinal, alternating series, disposed all around the branch Cellariafistulosa (Fig. 7)

Colony jointed or not; zooids uni- or multi-serial, all facing the same way 18

18 Colony in the form of a miniature tree with a tall, slender, cylindrical stem surmounted by a crown of

fiexible branches composed ofbiserially arranged zooids (Fig. 8). Each zooid with a conspicuous proximal

flexor muscle (seen through frontal membrane with transmitted light). A slender pedunculate avicularium

attached laterally to many zooids. Offshore only, on muddy bottoms (Kinetoskias) 19

Colony not as described , , , 20
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 11

Figure 8

Kinetoskias smitti (after Maturo and Schopf 1968

[Scale is approximate)).

19 Colony with one main stem, and few to many additional rootlets attaching it to the substratum. Branches

of crown forming a regular funnel shape. Avicularia of one size only, distinctly humpbacked, attached

at distal end of zooid (see Fig. 56) Kinetoskias arborescens

Colony with only a single stem, formed through the fusion of several rootlets, up to 5 em long. Crown

more or less bilobed. Avicularia of two sizes, slender, attached to lateral margins of zooids (see Fig. 57)

................................................................................. Kinetoskias smitti

20 Branches of colony uniserial (i.e., in the form of chains of zooids); colonies generally very small « 1 em

in height) 21

Branches of colony bi- or multi-serial; colonies generally (but not necessarily) exceeding 1 em in height

.............................................................................................. 24
12 NOAA Technical Report NMFS 99 _

21 Zooids horn-shaped, deeper than wide, with an elongate oval-rectangular membranous area which lacks

surrounding spines; new chains of zooids initiated frontally, just below the membranous area; terminal

brood-chambers subglobular, bivalved (Scruparia) 22

Zooids club-shaped, not deeper than wide, with either an oval, spine-surrounded membranous area or a

sub-circular operculate orifice; new chains of zooids never initiated frontally; terminal brood chambers

(ovicells) globular, not bivalved 23

22 Erect chains of zooids arise from creeping lines of zooids; zooids slender in profile, with the membranous

area nearly parallel to the zooidal axis (see Fig. 31) Scruparia ambigua

Erect chains of zooids arise from creeping stolons; zooids more strongly horn-shaped, with the membranous

area clearly oblique to the zooidal axis (see Fig. 32) Scruparia chelata

23 Stems jointed; branching dichotomous; zooids with a membranous frontal area surrounded by a number

of spines; ovicells imperforate (see Fig. 58) Bugulellafragilis

Stems unjointed; branching irregular; zooids ascophoran (i.e., with the membranous area replaced by

a subcircular, operculate orifice), the orifice sinusoid; ovicells perforate (see Fig. 68) Haplota clavata

24 Some or all the zooids bearing a loosely articulated, mobile, "bird's head" avicularium 30

"Bird's head" avicularium absent (though sessile avicularia may occur); zooids may support a frontal

shield (scutum) overarching the membranous area frontally and/or a vibraculum with a mobile, whip-like

seta basally 25

25 Colonies white and slender; zooids without scutum, without vibracula, and without either lateral or

frontal avicularia (see Fig. 42) Tricellaria peachii

Colonies of variable form; zooids with scutum and/or vibracula, and with lateral and (frequently) frontal

avicularia 26

26 Branches bi- to multi-serial, apparently unjointed; large vibracula present, each with a serrate seta; basal

surface of the bearing zooid obscured by the large, obliquely oriented, vibracular chambers; rhizoids from

the vibracular chambers meeting centrally, passing as a ridge down the branches and proximally forming

bundles of anchoring rootlets; no frontal scutum (see Fig. 38) Caberea ellisii

Branches biserial, clearly jointed; vibracula absent or, if present, small, with smooth seta, the chamber

covering only a small proportion of the basal surface; anchoring rhizoids fewer, not forming bundles; frontal

scutum well developed, or small and finger-like, sometimes absent 27

27 Vibracula absent. Usually three to five zooids in the internode (up to nine distally). Joints not traversing

the membranous area of any zooid at the bifurcation 28

Vibracula present or absent. Usually five to 12 zooids in the internode (up to 20 distally). Joints traversing the membranous area of the two outside zooids at the bifurcation 29

28 Internodes most frequently of three zooids, up to five or seven in distal parts of colony. Scutum roundedtriangular, covering half or less of frontal membrane (see Fig. 44) Tricellaria ternata

Internodes most frequently of five to nine zooids, sometimes more in distal parts of colony. Scutum oval,

covering most of frontal membrane, except for operculum (see Fig. 43) Tricellaria gracilis

29 Scutum rounded, triangular or oval; small, covering no more than half area of frontal membrane (see

Fig. 40) Scrupocellaria scabra

Scutum rounded rectangular to elongate oval, distal edge often straight; very large, obscuring most of

frontal membrane and extending distally to hide part of ovicell (see Fig. 41) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scrupocellaria scabra var. paenulata
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 13

30 Colony a white, feathery tuft; branches biserial. Zooids slender proximally, much wider above; the membranous area oval, less than one half length of zooid, fringed by a series of long, curved spines; ovicell

globular, attached by a narrow peduncle on the inner border of the membranous area .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bicellariella ciliata (Fig. 9)

Colony generally yellowish, buff or gray; branches bi- to multi-serial. Zooids only slightly narrower proximally; membranous area elongated, generally occupying most of frontal surface of zooid; ovicell crescentic to globular, terminal on and orthogonal to the proximal zooid (slightly oblique in one species) 31

Figure 9

Bicellariella ciliata. Scale: 5 mm.

31 Seen basally, the cross walls between zooids pointed distally, forked proximally; except in zooids near

the ancestrula, marginal spines confined to distal angles; avicularia attached about midway or distally

on the zooid wall (Bugula) 32

Seen basally, the cross walls between zooids are orthogonal to the zooidal axis; a spine or spines may

be confined to the distal zooid angles or present on the lateral margins as well; avicularia attached to the

frontal zooid wall, proximal to the membranous area (Dendrobeania) 36

32 Branches with zooids in more than two series (three to six, occasionally more), with avicularia borne only

by marginal zooids; ovicells hemispherical (see Fig. 48) Bugula simplex

Branches with biserially arranged zooids (or if multiserial distally then avicularia borne by both inner

and marginal zooids); ovicells saucer-shaped, spherical or subglobular 33

33 Branches becoming multiserial distally; if biserial, when viewed from the basal side (except perhaps near

the origin of the colony), at least one enclosed zooid below the axil of a bifurcation 34

No enclosed zooid at bifurcations 35

34 Outer distal angle of zooids bearing three spines, inner angle with at least two; most commonly at least

two enclosed zooids below the axil of a bifurcation (perhaps becoming multiserial distally) (see Fig. 46)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bugula fulva

Outer distal angle of zooids bearing two spines, inner angle with one; most commonly with one enclosed

zooid below the axil of a bifurcation (see Fig. 49) Bugula stolonifera

35 Outer distal angle of zooids bearing two to three spines, inner angle with two; at least some avicularia

longer than the width of the zooid at the point of attachment; ovicells hemispherical, orthogonal to the

branch axis (see Fig. 47) Bugula harmsworthi
14 NOAA Technical Report NMFS 99 _

Colony tufted, with spiralling branches. Outer distal angle of zooids bearing a single spine, with beside

it a pointed tooth which is usually bent somewhat over the membranous area; inner angle with one spine.

Avicularia small, their length less than the width of the zooid at the point of attachment; ovicells slightly

oblique to the branch axis Bugula turrita (Fig. 10)

Figure 10

Bugula turrita. Scale: 20 mm.

.'36 Branches of colony with zooids in two series .'37

Branches of colony with zooids in three or more series .'38

37 Distal angle of zooid bearing one distally directed projection (very rarely there may be one lateral spine),

inner angle with or without spine; avicularia short and broad, about as long as high (see Fig. 55) ...

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrobeania decorata

Usually two spines on each distal angle, occasionally only one on the inner; avicularia almost twice as

long as high, of two sizes Dendrobeaniafruticosa var. quadridentata

38 Avicularia of one type, present only on zooids of the inner series (see Fig. 53) Dendrobeania murrayana

Avicularia of two types: large on zooids of the marginal series, small on inner zooids (the two kinds also

differing somewhat in shape) 39

39 Marginal zooids bearing one spine on the outer distal angle, with at most one spine proximal to it (see

Fig. 52) Dendrobeania fruticosa

Marginal zooids bearing one spine on the outer distal angle and at least two or three lateral spines along

the outside of the membranous area 40
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 15

40 Marginal zooids with five to ten spines along the outside of the membranous area (see Fig. 54) .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrobeania pseudomuTTayana

Marginal zooids with two to three spines along the outside of the membranous area .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrobeania pseudomuTTayana var. fessa

41 Colony delicate, comprising an encrusting uniserial chain of zooids, giving rise to erect uniserial shoots;

each zooid budded from the disto-basal surface of its predecessor (see Fig. 68) Haplota clavata

Colony robust, multiserial 42

42 Colony form variable, but generally with short, irregular and rather nodular branches; often more nearly

mamillate than branching. Zooids in ordered series at growing tips, elsewhere with random orientation.

Large spatulate avicularia present Celleporina surcularis (Fig. 11)

Colony freely branching, neither mamillate nor nodular. Zooids in ordered longitudinal series 43

Figure 11

Celleporina surcularis. Scale: 10 mm.

Figure 12

Pseudoflustra solida. Scale: 5 mm.

43 Branches slender, or broadly lobate, but always bilaminate and flat 44

Branches distinctly cylindrical ................................................................... 48

44 Colony attached to the substratum by bundles of brown, chitinized kenozooidal tubes. Each zooid with

a large, triangular avicularium immediately proximal to orifice Pseudoflustra solida (Fig. 12)

Colony attached to substratum by an encrusting calcareous base 45
16 NOAA Technical Report NMFS 99 _

45 Each zooid with a prominent, bulbous avicularium medio-proximal to the orifice. Frontal surfaces of zooids

smooth, without obviou; pores. Colony generally with slender branches Cystisella saccata (Fig. 13)

Avicularia, when visible, lateral to orifice, usually transversely oriented 46

Figure 13

Cystisella saccata. Scale: 10 mm.

Figure 14

Rhamphos/omella cos/a/a. Scale: 5 mm.

46 Zooid orifice widely open, flanked by lateral peristomial flaps, and with a large, transversely oriented

avicularium with hooked tip. Ovicells prominent, porous. Colony lobed, little branched .

· Rhamphostomella costata (Fig. 14)

Zooid orifice generally deeply immersed. Avicularium small, sometimes inconspicuous. Ovicell indistinct,

imperforate, or with a single small pore 47

47 Aperture of developed zooids more or less pear-shaped, with a transversely oriented, lateral oral avicularium.

Ovicell a small cap-like structure with a single frontal pore. Colony slender, richly branched .

· Posterula sarsi (Fig. 15)

Aperture of developed zooids with a slit-like proximal sinus. Avicularium oriented proximo-laterally. Ovicell

imperforate, immersed, and rarely visible. Colony typically with broad, lobate branches .

· Ragionula rosacea (Fig. 16)
__________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 17

Figure 15

Posterula sarsi. Scale: 10 mm.

Figure 16

Ragionula rosaaa. Scale: 5 mm.

Figure 17

Porelloides laevis. Scale: 5 mm.

48 Zooids opening on only two-thirds of the periphery of the branch; remaining part constituting a defined

basal surface formed of large kenozooids. Zooids with marginal pores only. Colony with tapered, cylindrical branches Porelloides laevis (Fig. 17)

Zooids opening around whole periphery of branch, which is smoothly cylindrical and densely porous 49
18 NOAA Technical Report NMFS 99 _

49 Avicularia interzooidal, the rostrum as large as the zooid orifice. Colony branches dividing irregularly

..................................................................... Myriapora coarctata (Fig. 18)

Avicularia adventitious, small; situated distolateral to zooid orifice, single or paired. Colony branches

regularly dichotomous Myriapora subgracile (Fig. 19)

Figure 18

Myriapora coarclala. Scale: 10 mm.

Figure 19

Myriaj;ora subgracile. Scale: 10 mm.
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 19

Annotated Systematic List _

Each species entry is accorded a single literature citation,

which will give the accepted synonymy for that species,

together with a complete description and additional

illustrations.

Class STENOLAEMATA

Calcified bryozoans with tubular zooids. The circular aperture of the zooid is covered by a terminal membrane, with

the orifice, through which the lophophore is everted, at its

centre. Calcified wall with pseudopores: uncalcified spots

covered externally by a thickened area of cuticle, and

plugged internally by a special cell. Polypide enclosed in

a membranous sac, dividing the body cavity of the zooid

into two parts. A single living order, exclusively marine,

and usually comprising about 10% of any bryozoan fauna.

Order CYCLOSTOMATA

Heavily calcified tubular bryozoans, with characteristic

speckled appearance imparted by densely spaced pseudopores. Lophophore eversion achieved by redistribution of

internal coelomic pressure. Polymorphism limited, including autozooids, gonozooids and kenozooids; gonozooids

typically large and swollen, and important in taxonomy.

Reproduction, uniquely, involves repeated division of the

primary embryo to produce numerous secondary and tertiary embryos.

Family CRISIIDAE

Colony slender, delicate, jointed, branching dichotomously; branches consisting of internodes with zooids in

uniserial or biserial arrangements, linked by chitinized

tubular nodes. Autozooids elongate, tubular, with apertures all facing in the same direction, usually surmounting a projecting peristome. Gonozooid pyriform, replacing an autozooid, its aperture (ooeciostome) typically a

slender, flared or reflected funnel. Spines may be present.

Colony attached to substratum by slender, kenozooidal

rhizoids.

Crisia cribraria Stimpson, 1853-[Osburn 1912:215, pI.

18, figs. 7, 7a-b) Fig. 20

Colony developing dense, bushy tufts, up to 3 cm high.

Internodes stout, long and flat, commonly with 18-20

zooids, but ranging from 10 to 26. Zooids 0.35-0.5 mm

long; projecting peristome of zooids very short, 0.007-

0.009 mm diameter. Gonozooid occupying position 5 or

6 in the internode (range from position 3 to 9), elongate,

conspicuously swollen distally; ooeciostome a short tube,

facing directly distally, flared at its tip, with a transversely

elongate orifice.

Figure 20

Crisia cribraria. Scale: 0.5 mm.

Sublittoral. Recorded by Osburn (1912) from Woods

Hole, which he considered to be the southern limit of its

range. Nothing further seems to be known of its distribution or ecology.

Crisia sp.-[Crisia eburnea: Osburn 1912: 215, pI. 18,

figs. 6, 6a-b] Fig. 21

Colony up to 4 cm high. Internodes short, incurved,

commonly with 5-7 zooids, but ranging from 4 to 10; free

distal portion of zooid turned forward at right angle to

branch axis. Joints yellowish, to dark brown in proximal

parts of colony. Zooids 0.32-0.5 mm long, peristomes

0.006-0.008 mm diameter. Gonozooid occupying position

2 or 3 in the internode; ooeciostome short, tapered with

slightly convex proximal edge, its orifice transversely oval,

directed obliquely forwards. On algae, hydroids, stones,

shells, pier piles; common and abundant in shallow, coastal

waters.

North American records of Crisia eburnea (L.) perhaps

refer to a different species from that widely distributed

around northwest European coasts. Osburn's (1912) illustrations show a more slender species, with more prominent zooid peristomes, than the European one, which has

rather short internodes with the zooid peristomes scarcely

projecting. Crisia "eburnea" has been reported along the

eastern American seaboard, from North Carolina to

Labrador, but all of these records require reassessment.

Osburn (1912) reported a third species of Crisia for

Woods Hole, which he assigned doubtfully to the northwest
20 NOAA Technical Report NMFS 99 _

Figure 21

Crisia sp. (after Osburn 1912 [Scale is approximate]).

European species C. denticulata (Lamarck). Winston (1982)

suggested that Osburn's material may have been C. elongata Milne Edwards, which at Woods Hole would have

been beyond the northern limit of its known range. Further material of this species is required before its identity

can be established.

Family TUBULIPORIDAE

Colony adnate, creeping or lobate, or semierect or erect,

with elongate dichotomously dividing branches. Erect

species always attached to the substratum by an encrusting

portion of the colony; autozooids always opening on a

defined frontal surface. Tubular distal portions of zooids

either independent, or fused in connate rows. Gonozooid

occupying main axis of colony, extending laterally between

autozooid rows, its opening (ooeciostome) surmounti.ng a

slender tube, often flared or hooded distally.

Idmidronea atlantica (Forbes inJohnston, 1847)-[Hayward and Ryland 1985:90, fig. 31] Fig. 22

The colony is built of slender, regularly dichotomizing

branches, developing a two-dimensional fan shape. The

slender distal portions of the zooids are fused in connate

rows of three to five, deflected alternately to left and right

Figure 22

Idmidronea atlantica. Scale: 1 mm.

of the branch axis. The gonozooid is elongate and proximally tapered, with a slender, recurved ooeciostome.

This species is widely distributed in the northeast Atlantic, from Norway to Angola, and throughout the Mediterranean. It occurs offshore on coarse shell gravels, or on

rocky substrata, and in the Mediterranean is common in

a wide range of circalittoral habitats (Harmelin 1973,

1976). I. atlantica has been reported, reliably, from the Gulf

of Mexico (Harmelin 1976) and it is possible that it ranges

along the northeast coasts ofN. America, on the outer continental shelf. Kluge (1962, 1975) cites many Arctic

records, including the Canadian Arctic and the Gulf ofthe

St. Lawrence, but these need to be re-examined in the light

of recent systematic reviews (e.g., Harmelin 1976).

Family ANNECTOCYMIDAE

Colony adnate, lobate, or erect and irregularly branching.

Erect parts of colony with zooids arranged in whorls, opening around entire periphery. Zooids in alternating sequence, or in regular transverse rows, but the distal portions free and unfused, except at their bases. Gonozooid

elongate but not laterally lobed.

Entalophoroecia deflexa (Couch, 1842)-[Hayward and
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 21

Figure 23

Entalophoroecia deflexa. Scale: 1 mm.

Ryland 1985: 113, figs. 7C, 39]. Fig. 23

This extraordinarily plastic species has been reported

from throughout the North Atlantic region. In the northeast Atlantic it has been accorded a number of distinct

generic and specific names (see Hayward and Ryland 1985,

for synonymy). Some colonies are entirely adnate, developing open, regularly branching networks on flat substrata.

Unlike the superficially similar Annectocyma major Gohnston), these are developed by repeated dichotomy, and

adventitious branching does not occur. Zooid peristomes

are quincuncially arranged, with diameters of 0.13-0.20

mm. The gonozooid is steeply convex, pyriform in shape,

with a slender, unflared ooeciostome. Erect colonies may

be branched or unbranched, but are typically clubbed

distally, and the gonozooid may occupy most of the distal

regIOn.

E. deflexa is widespread in the northeast Atlantic from

Arctic Norway to the Mediterranean. It ranges from the

shallow sublittoral, offshore to at least 100 m, on stones,

shells, hydroids, algae, and other bryozoans. Northwest

Atlantic records are difficult to assess, but reports of

"Entalophora clavata" from the Gulf of St. Lawrence possibly

refer to this species, and it may prove to be generally

distributed off the northeast coasts of North America.

Family HORNERIDAE

Colony robust, richly branched, attached by a thick, encrusting base, thickening continuously through frontal

budding of zooids and secondary calcification, so that old

colonies may measure 10 mm or more at the base. Zooids

budded from the basal side of each branch, all opening

on a defined frontal surface, their peristomes distinct at

branch apices, immersed in thick calcification in oldest

parts of colony. Gonozooid grossly inflated, situated on

basal surface of branch.

Hornera lichenoides (Linnaeus, 1758)-[Hayward and

Ryland 1985:116, figs. 40, 41]. Fig. 24

Young colonies in the form of two-dimensional fans,

branching dichotomously to form stout three-dimensional,

reticulate structures up to 60 mm length. Base in old colonies often more than 10 mm diameter, branches tapering distally. Autozooids arranged in up to five alternating

longitudinal series at branch tips, increasing to 20 or more

series by continued budding of new series at all levels of

the colony. Apertures facing outwards, away from colony

axis, on a defined frontal surface; becoming immersed in

thick, granular calcification, with longitudinal folds and

furrows, and numerous large pores. Basal surfaces of

branch granular to nodular, with similar ridges, furrows

and pores. Gonozooid egg-shaped, up to 3 mm long,

densely perforated, with coarsely ridged surface.

H. lichenoides is widespread in Arctic waters from Siberia

to Greenland. In the eastern North Atlantic it ranges

southwards to the Western Isles of Scotland, where it occurs on coarse gravels, in depths greater than 50 m. It has

been reported on North American coasts as far south as

the Gulf of St. Lawrence, and may range further south on

the outer continental shelf.

Family STEGOHORNERIDAE

Colony stout, thickly calcified, attached by a thickened,

encrusting base, developing a smooth, non-porous outer

layer of calcification in late ontogeny. Autozooids budded

from axis of branch, apertures all opening on a defined

frontal surface, becoming immersed in calcification as

colony thickens; new series of autozooids budded at all

levels of colony. Gonozooid inflated, on basal surface of

branch, usually close to the dichotomy.

Stegohornera violacea (M. Sars, 1863)-[Stigmatoechos violacea: Hayward and Ryland 1985:120, fig. 42] Fig. 25

Young colonies fan-like, two-dimensional; developing a

complexly reticulate, three-dimensional form, up to 40 mm
22 NOAA Technical Report NMFS 99 _

Figure 24

Hornera lichenoides. (Left) Frontal view. Scale: 1 mm.

(Right) Basal view. Scale: 0.5 mm.

Figure 25

Stegohornera violacea. Scale: 1 mm.

high; pale violet when living, its surface appearing smooth

to the unaided eye. Autozooids arranged in up to five alternating, longitudinal series at branch tips, increasing by

continued budding at all levels of colony. Apertures all

facing outwards, away from colony axis, becoming immersed in smooth, thick calcification with a few scattered

pores on younger areas; basal surface smooth, finely punctate. Gonozooid ovoid or irregular, surface dimpled with

numerous small pores.

This species has a limited distribution in the northeast

Atlantic, from the Barents Sea and Spitzbergen, to the

southwest British Isles. It occurs offshore on coarse gravels,

and does not seem to extend into the shallow waters ofthe

North Sea. S. violacea was reported from the Gulf of St.

Lawrence by]ullien and Calvet (1903), but the record has

still to be verified.

Class GYMNOLAEMATA

Calcified or uncalcified bryozoans with cylindrical or squat

zooids. Orifice closed by sphincter muscles, or by a hinged

operculum. Lophophore eversion achieved through deformation of part of body wall. Zooid polymorphism well

developed.

Order CTENOSTOMATA

Zooids cylindrical, bottle-shaped or flattened; always uncalcified. Orifice terminal or frontal, closed by muscular

contraction. Embryos usually brooded within parent zooid.
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 23

~-------------------------------,

Figure 26

Alcyonidium verrilli (after Osburn, 1912). Scale:

20 illill.

Figure 27

Nolella stipata. Scale: 0.5 illill.

tubular distal portion, and slender, often filiform proximal

portion; forming linear chains of autozooids, often grouped

in clumps to form a dense turf, or fused to form an upright,

branching tuft. No kenozooids.

L -----..J

Nolella stipata Gosse, 1855-[Hayward 1985:88, fig. 26]

Fig. 27

Autozooids packed together in dense clumps; cylindrical,

up to 3.0 mm long, 0.2 mm wide, opaque grey-brown,

Family NOLELLIDAE

Colony creeping or erect. Autozooids columnar, with erect,

Alcyonidium verrilli Osburn, 1912-[Osburn 1912:252,

pI. 28, figs. 75, 75a-b; pI. 31, figs. 92, 92a] Fig. 26

Colony forming a richly branched, shrubby growth up

to 40 cm high; branches cylindrical or flattened, 2 to 3 cm

in diameter, sometimes distinctly palmate or antlerlike;

reddish to greyish brown in color. Zooids with 16 tentacles.

This species has not been described or reported since

Osburn's (1912) detailed account of Woods Hole material.

He considered it to be common, subtidally, fI'om Long

Island Sound to Martha's Vineyard. Osburn (1912, 1933)

distinguished this species from Alcyonidium gelatinosum (L.),

which he considered to be rare in the Gulf of Maine. In

northwest European waters this name has been applied

erroneously to a number of erect, branching species of

Alcyonidium, and if Osburn was correct in supposing that

a second such species occurred along northeast American

coasts, then it requires a new specific name.

Colony encrusting, forming thin translucent sheets, or thick

fleshy patches; or erect, forming firm, fleshy lobes. Autozooids rectangular, with frontal orifice, often papillate;

typically contiguous, sometimes only loosely coherent,

multilayered in erect species. Simple kenozooids may be

present, conical or papillate, not spinose.

Family ALCYONIDIIDAE

Polymorphs limited to spines, stolons and simple kenozooids. Mostly marine; a few freshwater species.
24 NOAA Technical Report NMFS 99 _

Figure 28

Anguinella palmata. Scale: 1 mm.

annulate, coated with detrital particles. Proximal part of

autozooid tapered, stoloniform, linked to neighbouring

autozooids. Lophophore with 18-22 tentacles.

Occurs on a range of substrata, including shell, stone,

hydroids and other bryozoans, common in shallow coastal

waters. N. stipata is distributed in the eastern North Atlantic from western Norway to West Africa, and throughout

the Mediterranean. Maturo (1957) gives a range in the

western North Atlantic from Canada to Brazil.

Anguinella palmata Van Beneden, 1845-[Hayward

1985:92, fig. 29] Fig. 28

Colony forming dense, gray-brown, branching tufts up

to 20 em high; surface with a muddy texture, covered with

accretions of fine silt particles and, basally, with dense

epizooite assemblages. Autozooids elongate, the distal portion and orifice distinct, proximal parts unclear; 0.8-1.5

mm long, slightly incurved towards colony axis. Lophophore with 10 or 11 tentacles.

Occurs on stones, shells and small algae, and often attached to piers, dock pilings or pipes; intertidal, on sheltered muddy shores, tolerant of low, fluctuating salinity,

Figure 29

Amathia vidovici (after Prenant and Bobin 1956). Scale: 1 mm.

sometimes common in estuaries. Recorded from scattered

localities along the southern coasts of the North Sea, from

southwestern Britain, Zaire and Senegal. In the western

Atlantic it has been reported from Beaufort, North Carolina (Maturo 1957), and is abundant in the Indian River

area of Florida (Winston 1982).

Family VESICULARIIDAE

Colony creeping or erect, with bottle-shaped zooids arising directly from a slender, kenozooidal stolon; branching

frequently, forming a loose, dependent tuft, a stiff, dichotomously branching bush, or simply ramifying over the surface of the substratum. Autozooids single, in loose or dense

clumps, or in regular, doubled, comb-like rows, resembling

a set of pan-pipes.

Amathia vidovici (Heller, 1867)-[Winston 1982: 110,

fig. 9] Fig. 29

Colony forming stiff, dichotomously branching tufts, 2-3

em high, colorless to light brown. Stolon about 0.2 mm

wide, supporting small groups of four to eight pairs of

zooids, arranged in loose half-spirals around the stolon.

Zooid groups usually concentrated in distal portions of

internodes and not enveloping axes of dichotomies.

Lower shore and shallow subtidal, attached to a wide

variety of substrata. Distributed throughout North and

South Atlantic, also eastern Pacific, and Indian Ocean.

North as far as Gulf of Maine in western Atlantic.
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 25

Figure 30

Bowerbankia imbricata. Scale: 1 mm.

Bowerbankia imbricata (Adams, 1798)-[Hayward 1985:

139, fig. 47] Fig. 30

Colony creeping, or developing erect, densely bushy tufts

up to 5 cm high. Stolon rather thick, 0.3 mm wide; zooids

in irregular clumps of six to 20, or up to 50 in large colonies, not closely connate. Polypide with ten tentacles.

Intertidal and shallow sublittoral; attached to almost

any kind of substratum, but especially algae. Distributed

throughout the temperate North Atlantic; in the west,

reported from Woods Hole to Florida (Winston 1982).

B. gracilis Leidy has smaller, entirely creeping colonies, with

slender stolons «0.1 mm wide), and polypides with only

eight tentacles.

Order CHEILOSTOMATA

Zooids calcified to a greater or lesser extent; basically boxFigure 31

Scruparia ambigua. Scale: 0.5 mm.

like, with a frontal orifice, usually closed by a hinged operculum. Embryos brooded in globular brood chambersovicells, in modified gonozooids, or rarely in an unmodified

parent zooid. Spines often present around the zooid or its

orifice; numerous types of specialized heterozooids occur.

Suborder ANASCA

Zooids with at least part ofthe frontal surface membranous,

with the polypide usually visible through it; membrane may

be partly, or almost wholly underlain by a calcareous plate,

or overarched and partly hidden by rib-like spines, but is

always present.

Family SCRUPARIIDAE

Colony consisting of uniserial chains of slender, hornshaped zooids; the attached base may be a series of encrusting zooids, or a kenozooidal stolon. New chains

budded from the frontal or lateral surfaces of zooids, at

irregular intervals. Broad distal end of zooid with an oval

frontal membrane. Embryos brooded in keeled, bivalved

chambers borne by specialized reproductive zooids. No

spines or avicularia.

Scruparia ambigua (d'Orbigny, 1841)-[Ryland and

Hayward 1977:50, fig. 16] Fig. 31

The delicate, white, tufted colony is attached by an encrusting chain of autozooids. The frontal membrane is
26 NOAA Technical Report NMFS 99 _

--II

Figure 32

Scruparia chelala. Scale: 0.5 mm.

almost parallel with the basal wall of the zooid, which consequently has a slender, fusiform profile. The frontal membrane of the brooding zooid is longer than wide.

From low in the intertidal zone, offshore to about 50 m,

on stones, shells, algae, hydroids, bryozoans and decapods.

Almost ubiquitous, in temperate and tropical waters, but

absent from Arctic and Antarctic seas.

Scruparia ekelata (Linnaeus, 1758)-[Ryland and Hayward 1977:52, fig. 17] Fig. 32

The colony is similar to that of S. ambigua, but is attached

by a slender, encrusting, kenozooidal stolon. The plane

of the frontal membrane is oblique to that ofthe basal wall,

giving the zooid a clavate profile. The frontal membrane

of the reproductive zooid is transversely oval.

In similar habitats to S. ambigua, and often co-occurring.

Geographical distribution uncertain through prior confusion with the latter species, but known to be widely

distributed throughout the temperate North Atlantic.

Family EUCRATEIDAE

Zooids elongate, slender, tapered proximally; the frontal

membrane occupying the distal half ofthe zooid. No spines,

avicuiaria, or ovicells. Embryos brooded in membranous,

transparent sacs attached externally at the distal edge of

the operculum.

Figure 33

Eucralea loricala. Scale: 0.5 mm.

Eueratea lorieata (Linnaeus, 1758)-[Ryland and Hayward 1977:54, fig. 18] Fig. 33

The colony forms a tall, dense clump, commonly around

10 em, but up to 25 em tall, buff or light-brown, resembling a miniature poplar tree. Branches divide at acute

angles, new branches generally arising alternately to the

left and right of the main axis. Each branch comprises a

linear series of zooids, each pair joined by their basal surfaces (i.e., back to back), all with identical orientation. The

frontal membrane is a slender oval shape.

Distributed from the lower intertidal, offshore to at least

75 m, attached to a broad variety of substrata. Widely

distributed in the boreal North Atlantic, and in Arctic

waters. Osburn (1912) reported it as common at Woods

Hole, and (1933) considered Cape Cod to be its southernmost limit on the northeast American coast.

Family FLUSTRIDAE

Colony composed of nat, flexible fronds; broad or narrow,

dividing irregularly, attached by encrusting sheets of zooids

rather than rhizoids. Fronds unilaminar, or bilaminar, with

the two layers of zooids back to back. Zooids lightly calcified, rectangular, with the frontal surface usually entirely
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 27

Figure 34

Flustrafoliacea. Scale: 0.5 mm.

membranous, often bearing spines. Vicarious avicularia

present, replacing zooids in usual sequence. Ovicells small,

immersed within the surface of the frond.

Flustrafoliacea (Linnaeus, 1758)-[Ryland and Hayward

1977:76, fig. 27] Fig. 34

Colony forming a stiff, bushy clump, up to 20 cm high;

light gray-brown in color, with a crisp texture, and a

distinctive lemon-like smell when fresh. Fronds bilaminar.

The zooids are rectangular or linguiform, each with four

or five short, thick spines at the distal end. Avicularia occur at the division of zooid rows; about half the size of a

zooid, bearing a strongly chitinized, brown, semicircular

mandible, supported on a slightly raised rostrum. The

ovicell is an ovoid chamber, completely immersed in the

colony surface, appearing in frontal view as a short crescentic cap at the distal end of the brooding zooid.

Flustra Joliacea occurs sublittorally on coarse, current

swept bottoms, frequently in dense beds. In the northeast

Atlantic it ranges from the Barents Sea to Biscay, and is

abundant around the British Isles. It has been reported

from Greenland, and in the recent past has been recorded

from the Bay of Fundy (Powell and Crowell 1967).

Carbasea carbasea (Ellis and Solander, 1786)-[Ryland

and Hayward 1977:79, fig. 28] Fig. 35

The colony forms very lightly calcified, delicate, lobate

Figure 35

Carbasea carbasea. Scale: 0.5 mm.

fronds, which branch infrequently. Occurs in small tufts,

up to 12 cm high, light brown or colorless; the fronds are

unilaminar and translucent. Zooids elongate rectangular,

rounded distally, without spines. No avicularia or ovicells.

This boreal/Arctic species has a circumpolar distribution. In the eastern North Atlantic it ranges as far south

as Denmark, Germany and Brittany, and in the west extends southwards to the Gulf of St. Lawrence. It has not

been reported from the northeastern United States, but

may occur well offshore on hard grounds.
28 NOAA Technical Report NMFS 99 _

Figure 36

Securiflustra securifrons. Scale: 0.5 mm.

Securiflustra securifrons (Pallas, 1766)-[Ryland and

Hayward 1977:84, fig. 31] Fig. 36

The colony forms a dense tuft, up to 10 em high, composed of narrow, angular, wedge-shaped branches, bifurcating frequently; new branches also arise by budding from

the narrow kenozooids which form the edges of the fronds.

Bilaminar. Zooids very elongate, rectangular, each new

series arising from the lateral kenozooids. No spines.

Ovicell globular, completely immersed, with an aperture

distal to that of the zooid, shielded by two flattened processes developed from the lateral walls of the brooding

zooid. Avicularia quadrangular, about one-quarter length

of a zooid, interposed within a zooid series, not at the division of a series; mandible semicircular to subtriangular.

On hard substrata; distributed from low in the intertidal

to at least 100 m. A cold temperate species, with a circumboreal distribution, in the eastern North Atlantic S. securifrons ranges south to the coasts of Britain, Spain and

Portugal, and into the western Mediterranean. It has been

recorded from Labrador and Newfoundland, and may well

prove to extend south ofthe St. Lawrence on the outer continental shelf.

Family CELLARIIDAE

Colony forming often dense tufts; branching stems conFigure 37

Cellariafistulosa. Scale: 0.5 mm.

sisting of solid, white, cylindrical internodes, up to 1 em

long, linked by narrow, brown, chitinous joints; dividing

dichotomously. Zooids hexagonal or lozenge-shaped, in

alternating longitudinal series, opening around the whole

periphery of the branch. Frontal membrane visible in

freshly collected material as a glistening outer cuticle,

underlain by a concave, calcified shield, with the semicircular aperture (opesia) at the distal end ofthe zooid. Small,

interzooidal avicularia present in most North Atlantic

species. Ovicells completely immersed; internodes with

brooding zooids usually slightly swollen, ovicells recognised

by their frontal apertures.

Cellariafistulosa (Linnaeus, 1758)-[Ryland and Hayward 1977:122, figs. 56B, 58] Fig. 37

Forms diffuse clumps up to 6 em high; internodes commonly 4-5 mm long, 0.4-0.8 mm diameter, consisting of

five or six longitudinal series of zooids. Opesia semicircular,

with a distinct proximal cap, and two proximo-lateral denticles. Avicularium almost one-third length of autozooid,

the mandible crescentic, with a straight proximal edge,

oriented slightly transversely to long axis of internode.

Sublittoral, on hard substrata, attached to rocks, shells,

kelp holdfasts, and probably widely distributed over the

North Atlantic continental shelves. It was listed by Rogick

(1964) from Woods Hole, but there appears to be no
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 29

Figure 38

Caberea ellisii, frontal surface. Scale: 0.5 mm.

published account of North American material. Does not

extend into Arctic waters. The superficially similar Microporina articulata (Fabricius) occurs widely in the Arctic.

Family SCRUPOCELLARIIDAE

Colony forming an erect, dichotomously branching tuft,

2-3 cm high; jointed, or sometimes appearing unjointed;

attached by bundles of rhizoids. Branches unilaminar, the

zooids in two to four alternating longitudinal series, all

facing in one direction, so that the colony has defined frontal and basal surfaces. Zooids with an oval frontal membrane, frequently partly, or largely, hidden by a flattened

spine, the scutum; normal spines usually present at distal

end of each zooid. Heterozooids various, and often numerous: adventitious avicularia typically occur at frontal

and distolateral positions on feeding zooids, vibracula occur (sometimes absent) on basal surfaces ofthe colony, and

often at axils of dichotomies. Ovicells present, usually conspicuous, typically with large frontal fenestrae.

Figure 39

Cabcrca ellisii, basal surface. Scale: 0.5 mm.

Caberea ellisii (Fleming, 1814)-[Ryland and Hayward

1977:130, fig. 62] Figs. 38, 39

The colony forms a stiff, fan-shaped tuft, up to 3 cm

high, yellowish-brown in color, with rather stout branches

composed oftwo to four series of zooids; attached by thick

rhizoids originating from the basal surfaces of the zooids,

and passing down the mid-line of the colony branches as

a thickened, conspicuous ridge. Zooids short, rectangular,

the frontal surface almost entirely occupied by the frontal

membrane. No scutum; each zooid with two spines at its

outer distal corner and one on the inner. Small adventitious

avicularia occur, usually one on the outer distal corner of

each zooid, one or two on the frontal surface proximal to

the frontal membrane. Each zooid with a massive vibraculum on its basal surface, obliquely oriented so that the seta

extends outwards from the branch margin, and the groove

into which it is retracted trends from the outer distal to

the inner proximal corner of the zooid. Each rhizoid arises

• from the proximal edge of a vibraculum. Ovicell globose,

with an irregularly oval fenestra close to its aperture.

Offshore, attached to stones, shells, hydroids and other
30 NOAA Technical Report NMFS 99 _

Figure 40

Scrupocellaria scabra. Scale: 0.5 mm.

bryozoans. Widely distributed in the boreal North Atlantic, ranging south to Scotland in the east and to Massachusetts in the west. Osburn (1933) recorded it as common, offshore, on the New England coast.

Scrupocellaria scabra (van Beneden, 1848)-[Ryland and

Hayward 1977:136, fig. 65] Fig. 40

The colony forms a diffuse tuft, up to 2 em high, attached by long, slender rhizoids which grow independently towards the substratum, rather than forming a thick

bundle. Branches composed of two longitudinal series of

zooids; distinct joints present at the base of each new

branch, the plane of jointing passing through the opesia

of the outer, basalmost zooid of the branch. Zooids slender,

with the oval frontal membrane occupying just over half

the frontal length; scutum oval, crescentic or subtriangular,

covering about half of the frontal membrane; one or two

spines at the outer distal corner of each zooid, one at the

inne;. A prominent lateral avicularium present on each

zooid; obliquely oriented frontal avicularia may also occur. Small vibracula may be present or absent, on basal

Figure 41

Scrupoeellario. scabra var. paenulda. Scale: 0.5 mm.

surfaces of zooids, but do not occur in the axil of the

dichotomies. Ovicell wider than long, with an elongate,

transversely oval fenestra.

Offshore, attached to hydroids, bryozoans, stones, shells

and other hard substrata. A boreal!Arctic species with a

circumpolar distribution. In the eastern Atlantic it ranges

south as far as northeast Britain. Osburn (1933) recorded

it as common in the Gulf of Maine and considered it to

be at its southern limit at Cape Cod.

Scrupocellaria scabra var. paenulata Norman, 1903-

[Kluge 1962:380; 1975:458, fig. 240] Fig. 41

This variety is distinguished by its grossly enlarged

scutum, which covers the whole of the frontal membrane,

and obscures part ofthe ovicell. In other respects it is identical with S. scabra.

Arctic, with circumpolar distribution. Occurs commonly

north of the St. Lawrence; Osburn (1912) reported specimens from Woods Hole which he considered to represent

var. paenulata.

Tricellaria peachii (Busk, 1851)-[Ryland and Hayward

1977:145, fig. 70] Fig. 42

Colony forming a diffuse white tuft, up to 3 em high,
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 31

Figure 42

Tricellaria peachii. Scale: 0.5 mm.

attached by slender, independent rhizoids, mostly originating from basal parts of colony. Zooids in two longitudinal series; branches usually with seven to nine zooids

between bifurcations, narrowed and distinctly jointed

basally. Zooids elongate, tapered, with the oval frontal

membrane occupying up to two-thirds total frontal length.

A single short spine at the outer distal corner of each

zooid, none on the inner; scuta absent. No avicularia or

vibracula. Ovicell globose, with a large, semicircular, frontal fenestra.

Offshore, attached to a variety of substrata. Essentially

a boreal species, ranging from Spitzbergen to northern Britain in the eastern North Atlantic, and south to Massachusetts in the west. Osburn (1912) reported it as rare at

Woods Hole and (1933) regarded Cape Cod as its southern

limit.

Figure 43

Tricellaria gracilis. Scale: 0.5 mm.

Tricellaria gracilis (van Beneden, 1848)-[Kluge 1962:

375; 1975:452, fig. 236] Fig. 43

Branches biserial. Zooids elongate, almost parallel-sided;

usually five to nine per internode, but up to 16. Frontal

membrane occupying less than one-third total zooid length;

two spines at the outer distal corner of each zooid (one often

greatly enlarged), one at the inner distal corner. Scutum

oval, large, obscuring most of frontal membrane. Lateral

avicularium conspicuous. Ovicell with a narrow, transversely oval frontal fenestra.

Offshore, attached to a variety of substrata. Essentially

an Arctic species, with circumpolar distribution. Powell
32 NOAA Technical Report NMFS 99 _

Figure 44

Tricellaria ternata. Scale: 0.5 mm.

(1968) gives Cape Cod as its southern limit, but there seems

to be no published account of New England material.

Tricellaria ternata (Ellis and Solander, 1786)-[Ryland

and Hayward 1977:144, fig. 69] Fig. 44

Colony forming a diffuse, delicate, white tuft, up to 3

cm high, attached to the substratum by slender, independent rhizoids. Zooids in two longitudinal series, each internode typically comprising just three zooids, but up to

five or seven in the longer distal branches of the colony;

the basal joint of each branch long and conspicuous. Zooids

elongate and tapered, with the small, oval, frontal membrane usually occupying no more than one-third total frontal length. Three evenly spaced spines at the distal end of

the zooid, the middle one absent in ovicelled zooids; a

small, rounded or triangular, scutum covers about onehalf ofthe frontal membrane. A conspicuously large lateral

avicularium on each zooid; small frontal avicularia sporadically present. No basal heterozooids. Ovicellionger than

wide, with one or few small frontal pores.

Sublittoral, in shallow water, attached to algae, hydroids,

bryozoans, and hard substrata. A boreal!Arctic species,

widely distributed in the North Atlantic, ranging southFigure 45

Bicellariella ciliata. Scale: 0.5 mm.

wards to northeastern Britain, and on the American coast

to Massachusetts. Osburn (1933) reported it as common

offshore along the coast of Maine.

Family BICELLARIELLIDAE

Colony unjointed, unilaminar, with zooids in two or more

longitudinal series; bifurcating frequently; attached by

rhizoids. Zooids typically trumpet-shaped, with a slender

proximal portion, and an expanded, oval, distal portion,

flared outwards at an acute angle to branch axis. Spines,

avicularia and ovicells present.

Bicellariella ciliata (Linnaeus, 1758)-[Ryland and Hayward 1977: 146] Fig. 45

Colony forming a spreading, feathery, white tuft, up to

3 cm high. Branches biserial, the zooids in strictly alternating sequence, with the projecting, oval distal portion,

bearing the frontal membrane, giving a zigzag profile to

the branch. Four to nine long delicate spines at the distal

end of each zooid, curling inwards towards the branch axis

(often broken short by careless treatment), a single, similar
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 33

spine present on the proximal edge of the frontal membrane. Avicularia of the bird's-head type, with a serrated

upper beak, and a short, hooked mandible; a short stalk,

arising proximo-lateral to the frontal membrane. Ovicells

small, globose, on short stalks, attached to the inner lateral

edge of the frontal membrane.

Distributed from the lower intertidal, offshore, perhaps

to the edges of the continental shelves. Ranges widely in

the temperate North and South Atlantic.

Family BUGULIDAE

Colony unilaminar, branching, unjointed; biserial to multiserial, attached by rhizoids. Zooids lightly calcified, frontal surfaces almost entirely membranous, with or without

spines. Avicularia pedunculate, ofthe characteristic birdshead type. Ovicells usually present.

Genus BUGULA

Branches biserial to multiserial. Zooids elongate, tapered;

truncate distally; the proximal end forked in basal view.

Frontal membrane occupying almost all offrontal surface;

no operculum, the orifice instead closed by a sphincter.

Spines confined to distal ends of zooids. Avicularia attached

to lateral margins of zooids.

Bugulafulva Ryland, 1960-[Ryland and Hayward 1977:

160, fig. 77] Fig. 46

Colony forming a dense, fan-shaped, non-spiralled tuft,

up to 3 cm high, yellowish-brown in color. Branches

biserial. Zooids with three spines on outer distal corner,

two or three on inner distal corner; the most distal spine

on the outer, and the most proximal on the inner, often

greatly enlarged. Avicularia attached to margins of

zooids, just proximal to spines, slightly longer than the

width ofthe zooid, the beak smoothly downcurved. Ovicell

globose.

Occurs on the lower shore, and in the shallow sublittoral to about 70 m, attached to algae, bryozoans and hard

substrata. Distribution incompletely known, but occurs

widely in the eastern temperate North Atlantic, and in the

west from Maine to Brazil.

Bugula harmsworthi Waters, 1900-[Kluge 1962:340;

1975:409, fig. 213] Fig. 47

Colony small, spreading, cup-shaped, up to 2 em high.

Branches biserial. Zooids with two or three spines on outer

distal corner, one or two on inner distal corner. Avicularium attached midway along outer lateral margin of

zooid; elongate, almost halflength of zooid, with smoothly

downcurved rostrum. Ovicell hemispherical, widely open

frontally.

An Arctic species, reported from the Barents Sea to the

Davis Strait. It has been recorded offshore at Maine and

Figure 46

Bugulafulva. Scale: 0.5 mm.

Cape Cod (Osburn 1912, as B. cucullifera), but is probably

rare south of the St. Lawrence.

Bugula simplex Hincks, 1886-[Ryland and Hayward

1977:168, fig. 81] Fig. 48

Colony forming a broad, fan-shaped tuft, up to 3 em

high, light orange-brown to straw color. Branches with

three to six series of zooids, sometimes more. Zooids

with one spine on outer distal corner (rarely two) and

one on inner distal corner. Avicularia attached to zooids

of marginal series only, on the outer wall, about onequarter zooid length below the spines; as long as, or

slightly longer than, the width of a zooid, with the beak

smoothly downcurved. Ovicell hemispherical, the widely

open aperture occluded by a membranous vesicle during

brooding.

Distribution and ecology imperfectly known. In the

North Atlantic this species is frequently found associated

with docks, harbours and piers. Rogick (1964) records it

as abundant on pilings at Woods Hole. In much of the
34 NOAA Technical Report NMFS 99 _

Figure 47

Bugula harmsworthi (after

Kluge 1962 [scale approximate]).

Figure 48

Bugula simplex. Scale

0.5 mm.

Figure 49

Bugula stolonifera. Scale:

0.5 mm.

earlier literature dealing with the Woods Hole area, this

species was erroneously identified as B. flabellata.

Bugula stolonifera Ryland, 1960-[Ryland and Hayward

1977:170, fig. 82] Fig. 49

Colony forming a dense grey-brown tuft, up to 4 em

high. Branches biserial. Zooids with the outer distal corner drawn out as a spinous projection, often very long, and

with one, rarely two, accessory spines below it; inner distal

corner with a single spine. Avicularia attached to outer

margins of zooids, just proximal to spines, length about

equal to width of zooid, although the axillary zooid may

bear a very much smaller avicularium. Ovicell globose.

Distribution and ecology imperfectly known. In the

temperate North Atlantic it has been reported most often

from docks and harbours.

Bugula turrita (Desor, 1848)-[Winston 1982: 130, fig. 50]

Figs. 50, 51

Colony developing dense, bushy tufts, richly branched,

up to 10 cm high; main stem thickly chitinized, brown

colored, secondary branches disposed in a spiral around

main branch axes. Biserial; zooids tapered proximally, with
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 35

frontal membrane occupying about three-quarters total

length. One or two spines on outer distal corner, one on

inner distal corner. Avicularium attached midway along

outer lateral margin of zooid; small, less than the width

of the zooid, rostrum downcurved. Ovicell caducous, in

the form of a shallow, stalked cap, topping a swollen ooecial

vesicle, attached to inner side of distal margin of zooid and

inclined towards branch axis.

This species is abundant in shallow waters from Florida

to Massachusetts.

Genus DENDROBEANIA

Branches generally multiserial, less frequently biserial.

Zooids truncate both distally and proximally (not forked

proximally) in basal view. Frontal membrane occupying

almost all of frontal surface in median zooids, about onehalf in marginal zooids, the orifice closed by an operculum.

Spines present at distal ends of zooids, and typically also

along lateral margins. Avicularia attached frontally, proximal to the frontal membrane.

Dendrobeaniafruticosa (Packard, 1863)-[Kluge 1962:

331; 1975:398, fig. 206) Fig. 52

Colony a diffuse, delicate tuft, 2-3 em high, with narrow branches consisting of two to four zooid series.

Marginal zooids with one small spine on outer distal corner (rarely two), inner corner unarmed; median zooids with

two spines at each distal corner. Avicularia with downcurved beak; those of marginal and median zooids similar

in size, but the latter more slender and less domed than

former, Ovicell prominent, globular, smooth surfaced.

Distributed from the shallow subtidal to 300 m, or

deeper; attached to a variety of hard substrata, as well as

Figure 50

Bugula turrita. Scale: 0.5 mm.

Figure 51

Bugula turrita: development sequence ofthe ovicell. Scale: 0.25

mm.

5 6 7
36 NOAA Technical Report NMFS 99 _

Figure 52

Dendrobeaniafruticosa. Scale: 0.5 mm.

to bryozoans and hydroids. Boreal/Arctic, ranging south

to the northern North Sea, Greenland, Labrador and the

Gulf of St. Lawrence. Osburn (1933) reported it (as D. murrayana var. fruticosa) from the Maine coast.

Dendrobeaniafruticosa var. quadridentata (Loven, 1834)-

[Kluge 1962:333; 1975:400, fig. 207]

Branches with just two zooid series. Each zooid with two

short spines on outer distal corner, and two (rarely one)

on inner distal corner.

Occurs in similar habitats to D. fruticosa, but less frequently reported. Perhaps more strictly Arctic in distribution; recorded from Arctic U.S.S.R., Arctic Canada,

Labrador and Greenland.

Dendrobeania murrayana (Bean in Johnston, 1847)-

[Ryland and Hayward 1977:174, fig. 84] Fig. 53

Colony forming a dense, bushy tuft, up to 3 em high;

branches broad, flat, square-ended, with from four to 12

zooid series. Zooids with a single spine at each distal corner, and with two to seven lateral spines on each side.

Avicularia slender, with smoothly downcurved beak; those

of marginal zooids, when present, two to three times longer

Figure 53

Dendrobeania murrayana. Scale 0.5 mm.

than those of inner zooids. Ovicell prominent, globular,

with faint, radiating striations.

Shallow sublittoral, ranging offshore to the edges of the

continental shelves; on stones, shells and other bryozoans.

Boreal!Arctic, ranging south to the northern British Isles,

Greenland, and Cape Cod. Osburn (1933) reported it as

abundant on hard grounds along the Maine coast.

Dendrobeania pseudomurrayana Kluge, 1955-[Kluge

1962:328; 1975:394, fig. 203] Fig. 54

Colony forming a dense, bushy tuft, with broad, flat,

truncated branches, consisting of up to 20 zooid series.

Zooids with a single, stout spine at each distal corner, and

four to six stout, incurved spines along each lateral margin.

Avicularia of marginal zooids less than twice length ofthose

ofinner zooids, but proportionately very much deeper and

highly domed. Ovicell prominent, globular, with distinct

radiating sculpture.

Distributed from the shallow sublittoral to 200 m or

more, attached to shells, stones, hydroids and bryozoans.

Boreal/Arctic, distributed south to western Norway, Green-
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 37

Figure 54

Dendrobeania pseudomurrayana (after Kluge 1962 [scale is approximate)).

land, Arctic Canada, and perhaps to the Gulf of St.

Lawrence.

Dendrobeania pseudomurrayana var.fessa Kluge, 1955-

[Kluge 1962:330; 1975:397, fig. 205]

This variety was distinguished from the nominate species

by its fewer, more slender spines-only two or three on

each lateral margin ofthe zooid-and by its proportionately

larger marginal avicularia. Additionally, the avicularia of

the inner zooids are said to have thickly calcified stalks.

An Arctic/Boreal species occurring in similar habitats

to those of D. pseudomurrayana. Distributed southwards to

Norway and Greenland. Kluge (1962, 1975) considered

part of Osburn's (1912) material of D. murrayana from

Woods Hole to represent this form.

Dendrobeania decorata (Verrill, 1879)-[Bugula elongata:

Kluge 1962:345; 1975:415, fig. 218] Fig. 55

Colony forming bushy tufts 5-6 em high. Branches

biserial. Zooids long and slender; a single, short spinous

process on the outer distal corner, inner distal corner

rounded. Avicularia short, fat, with strongly curved rostrum; attached frontally, proximal to frontal membrane.

Ovicell globular, with distinct, bold, radial sculpture.

Offshore, attached to a variety of hard substrata.

Generally rare and poorly known; reported from Arctic

Figure 55

Dendrobeania decorata (after Kluge 1962 [scale is approximate)).

U.S.S.R., western Norway and the GulfofSt. Lawrence.

Reported from the Gulf of Maine (Verrill 1879C) but no

recent records. Verrill's material was redescribed by

Maturo and Schopf (1968), who showed it to be the same

species as that recorded from northeast Atlantic and Arctic

waters as Bugula elongata Nordgaard.

Kinetoskias arborescens Danielssen, 1868-[Kluge 1962:

349; 1975:419, fig. 220] Fig. 56

Colony consisting of a shallow, funnel-shaped crown of

dichotomously dividing branches, surmounting a slender,

elongate, monosiphonic stem up to 5 em high. Branches

formed from two, alternating, series of zooids; linked

basally by membranous kenozooids. Independent rhizoids

issue from basal parts of branches. Zooids linguiform, lightly calcified, without spines. Avicularia hump-backed, rarely

longer than width of zooid; sharply hooked, attached to

outer distal corner of zooids. Ovicell shallow, obliquely

oriented at distal end of zooid.

Offshore, on soft substrata, ranging to abyssal depths.

Boreal/Arctic; widely distributed in the North Atlantic
38 NOAA Technical Report NMFS 99 _

Figure 56

Kinetoskias arborescens. Scale: 1 mm.

region, southwards to western Norway, Greenland and the

Gulf of St. Lawrence. It was reported from Arctic Canada

by Powell (1968), but there are no recent records south

of Cape Sable, Nova Scotia.

Kinetoskias smitti Danielssen, 1868-[Kluge 1962:350;

1975:421, fig. 221] Fig. 57

Colony consisting of a shallow, bilobed, funnel-shaped

crown of dichotomously dividing branches, surmounting

a slender, polysiphonic stem up to 5 cm high. Branches

biserial; zooids with outer distal corner produced as a short

spinous process. Avicularia slender, attached midway along

lateral margin of zooids, variable in size, often very large

(exceeding twice width of zooid). Ovicells globular, attached to inner distal margin of zooids.

Offshore, in deep water and ranging to abyssal depths;

usually in unconsolidated sediments. Sporadically reported

throughout the boreal and Arctic North Atlantic region.

Described from the Gulf of Maine by Verrill (1875, as

Bugulaflexilis), but there are no recent records for the northeast American coasts.

Bugulella fragilis Verrill, 1879-[Maturo and Schopf

1968:36] Fig. 58

Figure 57

Kinetoskias smitti. Scale 1 rom.

Colony delicate, diffuse, consisting of uniserial branches,

dividing dichotomously at regular intervals. Zooids clavate,

with a slender proximal portion, and a rounded distal portion. Frontal membrane oval, occupying the entire fron:

tal surface of the distal portion of the zooid, surrounded

by eight to ten, short, incurved spines. A pedunculate

avicularium with semicircular mandible, situated at the

distal end of some autozooids. Ovicell spherical, as wide

as zooid, with faint ridges on surface.

This rare, little known species was described from deep

water off the New England coast (Verrill 1879a). There

have been no further records of it. Generally, species of

Bugulella are associated with abyssal habitats.

Suborder ASCOPHORA

Frontal surface of zooids completely calcified, apart from

the orifice; frontal wall smooth or rugose, with the boundaries of individual zooids distinct or more or less obscure,

entire or variously punctured by few to many pores.
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 39

Figure 58

Bugulella fragilis (after Maturo and Schopf 1968).

Family RHAMPHOSTOMELLIDAE

Colony typically encrusting; in some species developing

erect, folded or lobate sheets, but without an ordered,

regular form. Frontal shield of zooids umbonuloid, viz. an

overarched calcified lamina with a large distal aperture,

through which the operculum may be seen, set in a membranous frontal body wall. Aperture with a conspicuous

denticle on its proximal margin, and usually with pronounced lateral peristomial flaps; frontal shield with

marginal pores only, accentuated by intervening, medially

directed ribs. Avicularia present, typically associated with

aperture. Ovicell prominent, subspherical, opening high

above aperture.

Rhamphostomella costata Lorenz, 1886-[Kluge 1962:

537; 1975:654, fig. 375] Fig. 59

The colony is primarily encrusting, but often produces

erect lobes, broad or narrow, which may fold to give

bilaminar sheets, 2 to 3 em high. Zooids convex, each with

well marked ridges radiating from a prominent, conical,

suboral avicularium, with sharply hooked, laterally directed

rostrum. Orifice elongate oval, with narrow, anvil-shaped

tooth on lower margin. Ovicell wider than long, lightly

calcified, with closely spaced, irregular, frontal pores. Zooid

boundaries become indistinct in later ontogeny through

continued frontal thickening of calcification.

Figure 59

Rhamphostomella costala. Scale: 0.5 mm.

Offshore, on hard substrata. Essentially an Arctic

species, ranging south to Greenland, the Gulf of St.

Lawrence, and Woods Hole, where it was reported as not

uncommon by Osburn (1912).

Posterula sarsi (Smitt, 1868)-[Escharopsis sarsi: Kluge

1962:547; 1975:666, fig. 384] Fig. 60

Colony developing a richly branched, rigid, threedimensional structure 8 em or more high with an equivalent

horizontal spread, attached by an encrusting base.

Branches bilaminar, flat, 5-10 mm wide, curved or folded,

often concavo-convex in section, bifurcating irregularly,

often broadened, or distinctly palmate, distally. Zooids with

thick, coarsely granular frontal wall, with distinct marginal

pores. Orifice oval, notched proximally, with a small,

proximo-lateral avicularium on its rim. Ovicell hemispherical, with a small frontal foramen. The proximal region

of the colony thickens through continued frontal calcification and the orifices ofthe zooids become deeply immersed,

with the ovicell and avicularium becoming completely

hidden from view.
40 NOAA Technical Report NMFS 99 _

,-----------------------

Figure 60

Posterula sarsi. Scale: 0.5 illill.

On hard substrata, offshore, to at least 300 m. An Arctic,

circumpolar species, ranging south to northern Norway,

Greenland, the Gulf of St. Lawrence, and the coasts of

Alaska. Osburn (1933) recorded a single specimen from

the Gulf of Maine.

Family SMITTINIDAE

Frontal wall of zooids evenly perforated, or with marginal

pores only. Primary orifice with a distinct proximallyrula,

most often flanked by lateral condyles. Oral spines present or absent; peristome variously developed. Ovicell

prominent, with or without frontal pores, not closed by

zooidal operculum. Adventitious avicularia present, usually

associated with primary orifice of zooid.

Porelloides laevis (Fleming, 1828)-[Hayward and Ryland

1979:126, fig. 48] Fig. 61

---------------------------~

Figure 61

Porelloides laevis. Scale: 0.5 illill.

The colony forms a stout, rigid, dichotomously branching growth up to 4 em high, attached by a sheet of encrusting zooids. Branches slender, cylindrical, up to 2 mm

wide, sometimes slightly lobed at the tips prior to a

dichotomy. Zooids budded radially and at first opening

around whole periphery of branch; in proximal regions of

colony orifices of zooids on parts of the branch are occluded, defining a basal surface. Zooids tumid, smooth,

with inconspicuous marginal pores; primary orifice wider

than long, the proximal lyrula broad, but short and indistinct. No oral spines. Suboral avicularium with broad,

semielliptical mandible. Ovicell globular, without pores.

As calcification continues to thicken, boundaries between

zooids are obscured, and the primary orifice, avicularium

and ovicell are progressively immersed and hidden.

Offshore, on hard substrata. A boreal species, probably

widely distributed in the North Atlantic region. Recorded
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 41

Figure 62

Cystisella saccata: Zooids at growing edge.

Scale: 0.5 mm.

south to British Isles and western Mediterranean, and to

Gulf of St. Lawrence and Bay of Fundy.

Cystisella saccata (Busk, 1856)-[Porella saccata: Kluge

1962:462; 1975:561, fig. 309] Figs. 62, 63

Colony developing an open, irregularly branching form,

3-4 cm high, attached by an encrusting base. Branches

bilaminate, flat, up to 5 mm wide, often anastomosing.

Zooids elongate, convex, smooth surfaced and imperforate;

primary orifice terminal, almost perpendicular to frontal

plane, bell-shaped, with a convex proximal lip; no oral

spines. Avicularium with a fusiform chamber extending

entire frontal length of zooid; mandible terminal, situated

immediately proximal to zooid orifice, semicircular. Ovicell

prominent, lightly calcified, without pores, widely open

frontally. The boundaries between zooids become indistinct

as calcification thickens in late ontogeny; primary orifice,

avicularium, and ovicell are all progressively immersed.

Distributed from the shallow sublittoral, perhaps to the

edges of the continental shelves, attached to a variety of

Figure 63

Cystisella saccata: Zooids from older region of colony, three with

ovicells.

hard substrata. An Arctic species, ranging southwards to

Norway, Greenland, and the Gulf of St. Lawrence. This

species is inappropriately classified among the Smittinidae,

but its systematic affinities are presently unknown.

Family SCHIZOPORELLIDAE

Primary orifice of zooids orbicular or semiorbicular; bipartite, comprising a larger anter, and a smaller, rounded or

slit-like poster, usually termed the sinus. Frontal wall of

zooids evenly perforated, or with marginal pores only.

Avicularia present or absent. Ovicells present.

Pseudoflustra solida (Stimpson, 1854)-[Kluge 1962:441;

1975:535, fig. 292] Fig. 64

Colony 2 to 3 cm high, slender, branching irregularly;

narrowest proximally, attached by bundles of chitinous

rhizoids originating from lowest series of zooids. Branches

bilaminate, flat, 2 to 3 mm wide, broadening distally.

Zooids elongate, rectangular, with a glistening outer cuticle; flat, separated by raised ridges, the frontal walls with

large and conspicuous marginal pores. Primary orifice with
42 NOAA Technical Report NMFS 99 _

Figure 64

Pseudoflustra solida. Scale: 0.5 mm.

a broad, V-shaped sinus between prominent angular condyles; no oral spines. Avicularium suboral, with elongateoval, proximally directed mandible. Ovicell longer than

wide, partly immersed, with scattered frontal pores.

From the shallow sublittoral to at least 300 m, atJ:ached

to a wide range of hard substrata. Widely distributed in

Arctic and boreal North Atlantic waters, ranging southwards, in the west, to the Bay of Fundy and the Gulf of

Maine (Powell 1968). Four other species of Pseudoflustra are

known from Arctic/North Atlantic regions, and it is posFigure 65

Ragionula rosacea. (Left) Old, immersed zooids. (Right) Young

zooids. Scale: 0.5 mm.

sible that any or all of them may occur off the northeastern

coasts of the United States. Kluge (1962, 1975) describes

and illustrates all five species of Pseudoflustra.

Family STOMACHETOSELLIDAE

Colony encrusting, or developing erect, lobate growths.

Frontal wall of zooids thickly calcified, with marginal and

scattered frontal pores. Primary orifice orbicular, or with

a distinct sinus; typically immersed and hidden, the overlying secondary orifice with a narrow pseudosinus. Ovicell

deeply immersed, typically with a single frontal foramen.

Avicularia present or absent.

Ragionula rosacea (Busk, 1856)-[Hayward and Ryland

1979:212, fig. 901 Fig. 65

Colony with short, thick, broadly palmate, bilaminate

branches, irregularly lobed or divided; up to 2 em high,

attached by an encrusting base. Zooids with thickly

calcified, granular, frontal wall, with indistinct marginal

pores. Primary orifice with shallowly concave proximal

edge; visible only in newly budded zooids, rapidly immersed, the secondary orifice transversely oval, with a

slender, U-shaped, pseudosinus, surrounded by a raised

peristomial rim. A single adventitious avicularium, with

triangular mandible, proximo-lateral to secondary orifice.

Ovicell spherical, small, imperforate; immersed by secondary calcification and generally inconspicuous.
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 43

Figure 66

MYTiapora coaTc/a/a. Scale: 0.5 mm.

On hard substrata, offshore to at least 300 m. A boreal!

Arctic species ranging southwards to western Norway,

Scotland, Greenland and the Gulf of St. Lawrence. Osburn

(1933) reported a single specimen from the Gulf of Maine.

Family MYRIAPORIDAE

Colony erect or encrusting. When erect, typically with

slender, cylindrical, widely diverging branches, developing an open, three-dimensional form; attached by an encrusting base. Zooids opening around the whole periphery

of the branch; axis of branch comprising a core of slender

tubes which divide and diverge, passing between the

autozooids and opening on the surface of the colony as

angular pores, sealed by the external cuticle. Primary

orifice D-shaped, with a narrow proximal sinus. Zooid

boundaries not apparent; branch surface appears regularly

porous, with quincuncially arranged zooids, and sporadic,

small, adventitious avicularia. Ovicells completely immersed, appearing as simple swellings distal to the zooid

orifice. .

Myriapora coarctata (M. Sars, 1863)-[Leieschara coarctata:

Kluge 1962:507; 1975:616, fig. 353] Fig. 66

Figure 67

MYTiapora subgracile. Scale: 0.5 mm.

Colony up to 10 cm high. Branches 2-3 mm wide,

smoothly cylindrical in proximal parts of colony (where

zooid orifices are usually occluded by secondary calcification), distinctly moniliform distally; dividing dichotomously, or producing irregular lateral offsets. Avicularia randomly distributed between zooids, their mandibles as large

as, or larger than, the zooid orifice.

Offshore, attached to hard substrata. An Arctic species

with a circumpolar distribution. On northeast American

coasts reported as far south as Labrador and the Gulf of

St. Lawrence.

Myriapora subgracile (d'Orbigny, 1852)-[Leieschara subgracilis: Kluge 1962:510; 1975:619, fig. 355] Fig. 67

Colony up to, or just exceeding, 5 cm high. Branches

2-3 mm wide, smootWy cylindrical proximally, moniliform

or nodulated distally, dividing by regular dichotomy. Avicularia tiny, with semicircular mandibles; one or two

present close to the distal edge of the orifice of most zooids.

From the shallow sublittoral, offshore to at least 500 m,

on hard substrata. Essentially an Arctic, circumpolar
44 NOAA Technical Report NMFS 99 _

Figure 68

Haplota clavata. Scale: 0.5 mm.

species, ranging south to northern Norway, Greenland,

and the Gulf of St. Lawrence.

Family HIPPOTHOIDAE

Colony encrusting, or, rarely, erect. Frontal wall of zooids

thinly calcified, imperforate. Primary orifice rounded, with

a distinct proximal sinus. Avicularia generally absent, but

other types of polymorph frequent. Ovicell globular,

prominent, usually borne by morphologically distinctive

reproductive polymorphs.

Haplota clauata (Hincks, 1857)-[Hayward and Ryland

1979:256, fig. 110] Fig. 68

Figure 69

Celleporina surcularis. Scale: 0.5 mm.

Colony formed from upright, branching, uniserial chains

of zooids, arising from encrusting uniserial chains; 2-3 mm

high, branching irregularly. Zooids more or less clavate:

broadest distally, tapered proximally; each budded from

the disto-basal wall of its predecessor. Primary orifice orbicular, with a shallowly concave sinus. Ovicellate zooids

shorter than feeding zooids, the globular, terminal ovicell

with a frontal ridge and three or four large pores.

Probably widely distributed in the boreal North Atlantic, from the lower shore into the shallow subtidal. Typically associated with algae, hydroids and other erect bryozoans. Recorded from Denmark, northern British Isles,

Iceland, and the northeast coast ofAmerica from Cape Cod

to the Gulf of St. Lawrence. Osburn (1912) reported it as

rare at Woods Hole.

Family CELLEPORIDAE

Colony typically multilaminar, developing massive, nodular, or erect and freely branching forms. Zooids budded
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 45

frontally, without apparent orientation. Frontal walls with

marginal pores only. Adventitious and vicarious avicularia

present, often in abundance. Ovicells prominent, with

variable frontal perforation.

Celleporina surcularis (Packard, 1863)-[Cellepora surcutaris: Kluge 1962:553; 1975:673, fig. 388] Fig. 69

Colony forming a thick, dichotomously branching, threedimensional shape, 3-4 cm high; branches up to 5 mm

thick, rough surfaced, tapered distally. Zooids oval, convex. Primary orifice almost circular, with an indistinct

notch-like sinus; surrounded by a raised peristome, developed as a pronounced lip proximally, and bearing on each

side a small, columnar avicularium. Vicarious avicularia

randomly distributed among autozooids, of several types,

with semicircular, semielliptal, or distinctly spatulate mandibles. Ovicells prominent, spherical, with a depressed

frontal area, demarcated by a raised ridge, perforated by

small, round pores.

Offshore, on hard substrata. An Arctic species with circumpolar distribution, extending south to northern Norway, Iceland, Greenland, and the Gulf of St. Lawrence.

Several other species of erect Celleporidae, with similar

distribution, may perhaps occur in the north coast of the

American continental shelf (see Kluge 1962, 1975).

Citations

Abbott, M.B.

1973. Seasonal diversity and density in bryozoan populations of

Block Island Sound (New York, U.S.A.). In Living and fossil

Bryozoa (G.P. Larwood, ed.), p. 37-51. Acad. Press, London.

Harmelin, J.G.

1973. Morphological variations and ecology of the recent cyclostome

bryozoan 'Idmonea' atlantica from the Mediterranean. In Living

and fossil Bryozoa (G.P. Larwood, ed.), p. 95-106. Acad. Press,

London.

1976. Le sous-ordres des Tubuliporina (Bryozoaires Cyclostomes)

en Mediterranee, ecologie et systematique. Mem. Inst. Oceanogr. (Monaco) 10: 1-326.

Harmer, F.S.

1891. On the British species of Crisia. Q. J. Microsc. Sci. 32:

127-181.

Hayward, P.J.

1985. Ctenostome bryozoans. Synopses of the British Fauna (NS).

33. E.J. Brill/Backhuys, for the Linnean Society of London, and

the Estuarine and Brackish-water Sciences Association, London

and Leiden, 169 p.

Hayward, P.J., and J.S. Ryland.

1978. Bryozoa from the Bay of Biscay and western approaches.

J. Mar. BioI. Assoc. U.K. 58:143-159.

1979. British ascophoran bryozoans. Synopses of the British Fauna

(NS), 14. Academic Press, for the Linnean Society of London,

and the Estuarine and Brackish-water Sciences Association, London, 312 p.

1985. Cyclostome bryozoans. Synopses of the British Fauna (NS),

34. E.J. Brill/Backhuys, for the Linnean Society of London, and

the Estuarine and Brackish-water Sciences Association, London

and Leiden, 147 p.

Jullien, J., and L. Calvet.

1903. Bryozoaires provenant des campagnes de I'Hirondelle

(1886-1888). Result. Camp. Scient. Prince Albert I 23:1-188.

Kluge, G.A.

1962. Mshanki Severnykh Morei SSSR. Opred. Faune SSSR

76: 1-584.

1975. Bryozoa of the northern seas of the USSR. Amerind, New

Delhi Publishing Co., 1-711 p.

Maturo, F.J.S.

1957. A study of the Bryozoa of Beaufort, North Carolina, and

vicinity. J. Elisha Mitchell Sci. Soc. 73:11-68.

1968. The distributional pattern of the Bryozoa of the east coast

of the United States exclusive of New England. Atti Soc. Ital.

Sci. nat. Mus. Cir. Stor. nat. Milano 108:261-284.

Maturo, F.J.S., and T.J.M. Schopf.

1968. Ectoproct and entoproct type material: reexamination of

species from New England and Bermuda named by A.E. Verrill,

J.W. Dawson and E. Desor. Postilla, no. 20:1-95.

McKinney, F.K., and J .B.C. Jackson.

1989. Bryozoan evolution. Unwin Hyman, Boston, 1-238 p.

Osburn, R.C.

1912. The Bryozoa of the Woods Hole region. Fish. Bull., U.S.

30:205-266.

1933. Bryozoa of the Mount Desert region. BioI. SUTY. Mount

Desert region:3-97.

Powell, N.A.

1968. Bryozoa (Polyzoa) of Arctic Canada. J. Fish. Res. Board

Can. 25:2269-2320.

Powell, N.A., and G.D. Crowell.

1967. Studies on Bryozoa (Polyzoa) of the Bay of Fundy region.

II. Bryozoa from the intertidal zone of Minas Basin and Bay of

Fundy. Cahiers BioI. Mar. 8:331-347.

Rogick, M.D.

1964. Ectoprocta. In Keys for marine invertebrates ofthe Woods

Hole region (R.!. Smith, ed.), p. 167-182. Woods Hole Marine

Biological Laboratory.

Rogick, M.D., and H. Croasdale.

1949. Studies on marine Bryozoa. III: Woods Hole region

Bryozoa associated with algae. BioI. Bull. 96:32-69.

Ryland, J.S.

1967. Crisiidae (Polyzoa) from western Norway. Sarsia 29:

269-282.

1976. Physiology and ecology of marine bryozoans. Adv. Mar.

BioI. 14:285-443.

Ryland, J.S., and P.J. Hayward.

1977. British anascan bryozoans. Synopses of the British Fauna

(NS) 10. Academic Press for the Linnean Society of London,

London, 188 p.

Verrill, A.E.

1875. Brief contributions to zoology from the museum of Yale College, No. 32. Results of dredging expeditions ofT the New England

coast in 1874. Am. J. Sci. (3)9:411-415.

1879a. Op. cit., 42. Notice of recent additions to the marine fauna

of the eastern coast of North America, No.5, Polyzoa. Am. J.

Sci. (3)17:472-474.

1879b. Preliminary checklist of the marine Invertebrata of the

Atlantic coast, from Cape Cod to the Gulf of St. Lawrence.

Author's edition: Tuttle, Morehouse and Taylor, printers, New

Haven, Conn., 32 p.

1879c. Brief contributions to zoology from the museum of Yale College. 43. Notice of recent additions to the marine fauna of the

eastern coast of North America, No.6, Polyzoa, Echinodermata,

Anthozoa. Am. J. Sci. (3)18:52-54.

1879d. Notice of recent additions to the marine Invertebrata ofthe

northeastern coast of America, with descriptions of new genera

and species and critical remarks on others. Part 1: Annelida,
46 NOAA Technical Report NMFS 99 _

Gephyraea, Nemertina, Nematoda, Polyzoa, Tunicata, Mollusca,

Anthozoa, Echinodermata, Porifera. Proc. U.S. Nat. Mus.

2: 165-205.

Winston, J.E.

1982. Marine bryozoans (Ectoprocta) of the Indian River area

(Florida). Bull. Am. Mus. Nat. Hist. 173:99-176.

Systematic Index _

Alcyonidiidae 23

Alcyonidium gelatinosum 23

Alcyonidium verrilli 23

Amathia vidovici 24

Anasca 25

Anguinella palmata 24

Annectocyma major 21

Annectocymidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Ascophora 38

Bicellariella ciliata 32

Bicellariellidae 32

Bowerbankia gracilis 25

Bowerbankia imbricata 25

Bugula cucullifera 33

Bugula elongata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

Bugula jlabellata 34

Bugula jlexilis 38

Bugula fulva 33

Bugula harmsworthi 33

Bugula simplex 33

Bugula stolonifera 34

Bugula turrita . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

Bugulella fragilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 38

Bugulidae 33

Caberea ellisii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

Carbasea carbasea 27

Cellaria fistulosa 28

Cellariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

Cellepora surcularis 45

Celleporidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

Celleporina surcularis 45

Cheilostomata 25

Crisia cribraria . . . . . . . . . . . . . . . . .. 19

Crisia denticulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Crisia eburnea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 19

Crisia elongata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Crisiidae 19

Ctenostomata 22

Cyclostomata 19

Cystisella saccata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41

Dendrobeania decorata .. . . . . . . . . . . . . . . . . . . . . . . . . . . 37

Dendrobeania fruticosa 35

var. quadridentata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

Dendrobeania murrayana 36

var. fruticosa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

Dendrobeania pseudomurrayana 36

var. fessa 37

Entalophora clavata . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 21

Entalophoroecia deflexa. . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Escharopsis sarsi 39

Eucratea loricata 26

Eucrateidae 26

Flustra foliacea 27

Flustridae 26

Gymnolaemata 22

Haplota clavata 44

Hippothoidae 44

Hornera lichenoides 21

Horneridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Idmidronea atlantica 20

Kinetoskias arborescens 37

Kinetoskias smitti 38

Leieschara coarctata 43

Leieschara subgracilis 43

Microporina articulata . . . . . . . . . . . . . . . . . . . . . . . . . . .. 29

Myriapora coarctata 43

Myriapora subgracile 43

Myriaporidae 43

Nolella stipata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

olellidae 23

Porella saccata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 41

Porelloides laevis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

Posterula sarsi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

Pseudojlustra solida . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 41

Ragionula rosacea 42

Rhamphostomella costata 39

Rhamphostomellidae 39

Schizoporellidae 41

Scruparia ambigua 25

Scruparia chelata 26

Scrupariidae 25

Scrupocellaria scabra 30

var. paenulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Scrupocellariidae 29

Securiflustra securifrons 28

Smittinidae 40

Stenolaemata 19

Stegohorneridae 21

Stegohornera violacea 21

Stomachetosellidae 42

Tricellaria gracilis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31

Tricellaria peachii 30

Tricellaria ternata 32

Tubuliporidae 20

Vesiculariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Acknowledgments _

Preparation of this manual was supported in part by a grant

from the National Science Foundation to the Editorial
_________________ Ryland and Hayward: Erect Bryozoa of the Northeastern United States 47

Board of the "Marine Flora and Fauna of the Eastern

United States."

Preparation of manuals in the "Marine Flora and Fauna

of the Eastern United States" subseries is coordinated by

the following Board.

Coordinating Editor

Melbourne R. Carriker, College of Marine Studies, University of Delaware, Lewes, DE 19958.

Editorial Advisers

A. Ralph Cavaliere, Department of Biology, Gettysburg

College, Gettysburg, PA 17325.

Arthur G. Humes, Boston University Marine Program,

Marine Biological Laboratory, Woods Hole, MA 02543.

David L. Pawson, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Kenneth P. Sebens, Marine Science and Maritime Studies

Center, Northeastern University, East Point, MA 01908.

Ruth D. Turner, Museum ofComparative Zoology, Harvard University, Cambridge, MA 02138.

Robert T. Wike, Department of Botany, University of

Massachusetts, Amherst, MA 01002.

In addition to establishing the format for the "Marine

Flora and Fauna of the Eastern United States," the Board

invites systematists to collaborate in the preparation of

manuals, reviews manuscripts, and advises the Scientific

Editor of the National Marine Fisheries Service.

Coordinating Editor's Comments _

Publication of the "Marine Flora and Fauna of the Eastern United States" is most timely in view of the growing

universal emphasis on work in the marine environment and

the crucial need for precise and complete identification of

organisms related to this work. It is essential, if at all possible, that organisms be identified accurately to species.

Accurate scientific names of plants and animals unlock the

great quantities of biological information stored in libraries,

obviate duplication of research already done, and often

make possible prediction of attributes of organisms that

have been adequately studied.

John S. Ryland took his bachelor's degree in Natural

Sciences at Cambridge and his Ph.D. on settlement

behaviour in Bryozoa was with Dennis J. Crisp at the

University College of North Wales, Bangor. After an interlude in fisheries research he returned to academia and has

been at the University of Wales, Swansea since 1965. Bryozoan biology has always been an overriding interest, with

the book 'Bryozoans' published in 1970. Following a sabbatical in northern Australia and a long secondment to the

University of the South Pacific, Fiji, his area of interest

has extended to most of the colonial invertebrate groups,

and he has published on hydroids, zoanthids, entoprocts,

pterobranchs, and ascidians as well as Bryozoa.

Peter J. Hayward graduated from the University of

Reading in 1970, with a bachelor's degree in Zoology and

Geology. At the University of Wales, Swansea, he researched the population ecology of intertidal, epiphytic

bryozoans and the systematics and biogeography of Aegean

bryozoans for his Ph.D., which was awarded in 1974. Since

then he has published numerous papers on the Bryozoa,

and together with J. S. Ryland produced the four Linnean

Society Synopses of the British marine Bryozoa.
48 NOAA Technical Report NMFS 99 _

Published Manuals

Marine Flora and Fauna of the Northeastern United States

Annelida: Oligochaeta Cook, David G., and Ralph O. Brinkhurst

Protozoa: Ciliophora Borror, Arthur C.

Higher Plants of the Marine Fringe Moul, Edwin T.

Pycnogonida McCloskey, Lawrence R.

Crustacea: Stomatopoda Manning, Raymond B.

Crustacea: Decapoda Williams, Austin B.

Tardigrada Pollock, Leland W.

Cnidaria: Scyphozoa Larson, Ronald j.

Higher Fungi: Ascomycetes, Deuteromycetes, and Basidiomycetes

Cavaliere, A. R.

Copepoda: Harpacticoida Coull, Bruce C.

Sipuncula Cutler, Edward B.

Echinodermata: Holothuroidea Pawson, David L.

Copepoda: Lernaeopodidae and Sphyriidae Ho, Ju-Shey

Copepoda: Cyclopoids Parasitic on Fishes Ho, Ju-Shey

Crustacea: Branchiura Cressey, Roger F.

Protozoa: Sarcodina: Amoebae Bovee, Eugene C., and Thomas K. Sawyer

Crustacea: Cumacea Watling, Les

Arthropoda: Cirripedia Zullo, Victor A.

Cnidaria: Scleractinia Cairns, Stephen D.

Protozoa: Sarcodina: Benthic Foraminifera Todd, Ruth, and Doris Low

Turbellaria: Acoela and Nemertodermatida Bush, Louise F.

Lichens (Ascomycetes) of the Intertidal Region Taylor, Ronald M.

Echinodermata: Echinoidea Serafy, D. Keith, and F. Julian Fell

Echinodermata: Crinoidea Messing, Charles G., and John H. Dearborn

Erect Bryozoa Ryland, John s., and Peter j. Hayward

Marine Flora and Fauna of the Eastern United States

Cephalopoda Vecchione, Michael, Clyde F. E. Roper, and Michael j. Sweeney

Copepoda, Cyclopoida: Archinotodelphyidae, Notodephyidae, and

Ascidicolidae Dudley, Patricia L., and Paul L. Illg

NOAA Tech Rep.

NMFS Circular no.

374

378

384

386

387

389

394

397

398

399

403

405

406

409

413

419

423

425

438

439

440

446

NMFS no.

33

91

99

73

96

NTIS no.

COM 73 50670

73 50888

74 50019

74 50014

7450487

74 51194

PB 257 987

261 839

268036

268714

273 062

274999

280040

281 969

222 923

285 538

296 460

297 676

124 520

225 053

219 387

124 735

PC A03/MF AOI

PB 86 156 395

PB 89 189583